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Nutrient Requirements of Poultry: Ninth Revised
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ISBN: 0-309-59632-7, 176 pages, 8.5 x 11, (1994)
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Nutrient Requirements of Poultry
Ninth Revised Edition, 1994
Subcommittee on Poultry Nutrition
Committee on Animal Nutrition
Board on Agriculture
National Research Council
National Academy Press
Washington, D.C. 1994
Copyright © National Academy of Sciences. All rights reserved.
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ii
NATIONAL ACADEMY PRESS 2101 Constitution Avenue Washington, D.C. 20418
NOTICE: The project that is the subject of this report was approved by the Governing Board of the National Research Council, whose members are drawn from the councils of the National Academy of Sciences, the National Academy of Engineering, and the Institute of Medicine.
The members of the committee responsible for the report were chosen for their special competences and with regard for appropriate balance.
This report has been reviewed by a group other than the authors according to procedures approved by a Report Review Committee consisting of members of the National Academy of Sciences, the National Academy of Engineering, and the Institute of Medicine.
The National Academy of Sciences is a private, nonprofit, self-perpetuating society of distinguished scholars engaged in scientific and
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and technical matters. Dr. Bruce M. Alberts is president of the National Academy of Sciences.
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This study was supported by the Agricultural Research Service of the U.S. Department of Agriculture, under Agreement No.
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Library of Congress Cataloging-in-Publication Data
National Research Council (U.S.). Subcommittee on Poultry Nutrition.
Nutrient requirements of poultry / Subcommittee on Poultry
Nutrition, Committee on Animal Nutrition, Board on Agriculture,
National Research Council. — 9th rev. ed.
p. cm. — (Nutrient requirements of domestic animals)
Includes bibliographical references and index.
ISBN 0-309-04892-3
1. Poultry—Feeding and feeds. I. Title. II. Series: Nutrient requirements of domestic animals (Unnumbered)
SF494.N37 1994
636.5' 0852—dc20 94-3084
CIP
Copyright 1994 by the National Academy of Sciences. All rights reserved.
Any opinions, findings, conclusions, or recommendations expressed in this publication are those of the author(s) and do not necessarily
reflect the view of the organizations or agencies that provided support for this project.
Printed in the United States of America
First Printing, March 1994
Second Printing, April 1996
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iii
SUBCOMMITTEE ON POULTRY NUTRITION
JERRY L. SELL, Chair, Iowa State University
F. HOWARD KRATZER, University of California, Davis
J. DAVID LATSHAW, The Ohio State University
STEVEN L. LEESON, University of Guelph
EDWIN T. MORAN, Auburn University
CARL M. PARSONS, University of Illinois
PARK W. WALDROUP, University of Arkansas
COMMITTEE ON ANIMAL NUTRITION
HAROLD F. HINTZ, Chair, Cornell University
DONALD C. BEITZ, Iowa State University
GARY L. CROMWELL, University of Kentucky
DANNY G. FOX, Cornell University
ROGER W. HEMKEN, University of Kentucky
LAURIE M. LAWRENCE, University of Kentucky
LARRY P. MILLIGAN, University of Guelph, Canada
OLAV T. OFTEDAL, National Zoological Park, Washington, D.C.
JERRY L. SELL, Iowa State University
ROBERT P. WILSON, Mississippi State University
Staff
MARY I. POOS, Project Director
DENNIS BLACKWELL, Senior Project Assistant
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v
BOARD ON AGRICULTURE
THEODORE L. HULLAR, Chair, University of California, Davis
PHILIP H. ABELSON, American Association for the Advancement of Science, Washington, D.C.
JOHN M. ANTLE, Montana State University
DALE E. BAUMAN, Cornell University
WILLIAM B. DELAUDER, Delaware State University
SUSAN K. HARLANDER, Land O'Lakes, Inc., Minneapolis, Minnesota
PAUL W. JOHNSON, Natural Resources Consultant, Decorah, Iowa
T. KENT KIRK, U.S. Department of Agriculture, Forest Service, Madison, Wisconsin
JAMES R. MOSELEY, Jim Moseley Farms, Inc., Clark Hills, Indiana, and Purdue University
DONALD R. NIELSEN, University of California, Davis
NORMAN R. SCOTT, Cornell University
GEORGE E. SEIDEL, JR., Colorado State University
PATRICIA B. SWAN, Iowa State University
JOHN R. WELSER, The Upjohn Company, Kalamazoo, Michigan
FREDERIC WINTHROP, JR., The Trustees of Reservations, Beverly, Massachusetts
Staff
SUSAN OFFUTT, Executive Director
JAMES E. TAVARES, Associate Executive Director
CARLA CARLSON, Director of Communications
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PREFACE
vii
Preface
Formulation of balanced diets is fundamental to economical poultry production, and this process depends on a
knowledge of nutrient requirements of poultry and the nutritional attributes of nutrient sources. Thus, a compilation of
information on nutrient requirements and sources that can be used by feed formulators as a guideline is an important resource.
This ninth revised edition of the Nutrient Requirements of Poultry contains a reassessment of data used in the previous
edition and incorporates new information. The committee conducted an extensive review of the literature, and documentation
of most of this literature is included in this ninth edition. Note, however, that the review of literature was completed and the
nutrient requirements data compiled by the committee in September 1991.
The committee found that scientifically based knowledge about many nutrient requirements was incomplete.
Consequently, calculations and interpolations were necessary to derive estimated requirements for some nutrients. These
estimated requirements are identified in the requirements tables. In some instances, the committee decided that estimation of
the requirements was inappropriate and a question mark was used in the tables to indicate the absence of data.
Nutrient requirements given herein were derived, in most instances, from empirical observations of responses of poultry
to changes in dietary concentrations or intakes of specific nutrients. In some instances, nutritional models were used to
estimate amino acid requirements. Criteria used in establishing nutrient requirements included growth, reproduction, and feed
efficiency and, where possible, poultry health and quality of poultry products.
This report, as compared with previous editions, contains additional information on feedstuffs, including a description of
procedures used to determine metabolizable energy values and methods to estimate amino acid contents of feed ingredients.
A detailed discussion of dietary fat sources has been added, and the data presented on the nutrient composition of feedstuffs
have been expanded to include true metabolizable energy values and coefficients of true amino acid digestibility.
This ninth edition was prepared by the Subcommittee on Poultry Nutrition, which was appointed in 1989 under the
guidance of the Board on Agriculture's Committee on Animal Nutrition. The Committee on Animal Nutrition, the Board on
Agriculture, and several other experts reviewed the report. The subcommittee is grateful to these individuals for their efforts.
The subcommittee also thanks Roseanne Price for her editorial assistance and Mary Cochran and Ann Shuey of Iowa State
University for their secretarial assistance in preparing many drafts of the report.
JERRY L. SELL, Chair
Subcommittee on Poultry Nutrition
Copyright © National Academy of Sciences. All rights reserved.
PREFACE
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viii
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CONTENTS
ix
Contents
OVERVIEW
1
1.
COMPONENTS OF POULTRY DIETS
Energy
Carbohydrates
Protein and Amino Acids
Fats
Minerals
Vitamins
Water
Xanthophylls
Unidentified Growth Factors
Antimicrobials
3
3
8
9
11
13
15
15
17
18
18
2.
NUTRIENT REQUIREMENTS OF CHICKENS
Leghorn-Type Chickens
Meat-Type Chickens
19
19
26
3.
NUTRIENT REQUIREMENTS OF TURKEYS
Starting and Growing Turkeys
Turkey Breeders
35
35
39
4.
NUTRIENT REQUIREMENTS OF GEESE
40
5.
NUTRIENT REQUIREMENTS OF DUCKS
42
6.
NUTRIENT REQUIREMENTS OF RING-NECKED PHEASANTS, JAPANESE QUAIL, AND BOBWHITE QUAIL
Ring-Necked Pheasants
Japanese Quail
Bobwhite Quail
44
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44
44
45
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CONTENTS
x
7.
SIGNS OF NUTRITIONAL DEFICIENCIES IN CHICKENS AND TURKEYS
Protein and Amino Acid Deficiencies
Vitamin Deficiencies
Mineral Deficiencies
46
46
50
56
8.
TOXICITY OF CERTAIN INORGANIC ELEMENTS
58
9.
COMPOSITION OF FEEDSTUFFS USED IN POULTRY DIETS
Cereal Grains
Protein Supplements
Estimating the Amino Acid Composition of Feedstuffs
Characteristics of Dietary Fats
Macromineral Supplements
Mycotoxins
61
61
69
71
75
75
78
10.
STANDARD REFERENCE DIETS FOR CHICKS
80
A
B
C
APPENDIXES
Documentation of Nutrient Requirements
Estimating the Energy Value of Feed Ingredients
Conversion Factors
83
85
113
114
REFERENCES
115
AUTHORS
143
INDEX
145
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TABLES AND FIGURE
xi
Tables and Figure
Tables
1-1
1-2
1-3
2-1
2-2
2-3
2-4
2-5
2-6
2-7
2-8
3-1
3-2
3-3
4-1
4-2
Water Consumption by Chickens and Turkeys of Different Ages
Guidelines for Poultry for the Suitability of Water with Different Concentrations of Total Dissolved Solids
Xanthophyll and Lutein Content of Selected Ingredients
Nutrient Requirements of Immature Leghorn-Type Chickens as Percentages or Units per Kilogram of Diet
Body Weight and Feed and Water Consumption of Immature Leghorn-Type Chickens
Nutrient Requirements of Leghorn-Type Laying Hens as Percentages or Units per Kilogram of Diet
Estimates of Metabolizable Energy Required per Hen per Day by Chickens in Relation to Body Weight and
Egg Production
Typical Body Weights, Feed Requirements, and Energy Consumption of Broilers
Nutrient Requirements of Broilers as Percentages or Units per Kilogram of Diet
Nutrient Requirements of Meat-Type Hens for Breeding Purposes as Units per Hen per Day
Nutrient Requirements of Meat-Type Males for Breeding Purposes as Percentages or Units per Rooster per
Day
Nutrient Requirements of Turkeys as Percentages or Units per Kilogram of Diet
Growth Rate and Feed and Energy Consumption of Large-Type Turkeys
Body Weights and Feed Consumption of Large-Type Turkeys during the Holding and Breeding Periods
Nutrient Requirements of Geese as Percentages or Units per Kilogram of Diet
Approximate Body Weights and Feed Consumption of Commercially Reared Male and Female Geese to 10
Weeks of Age
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16
17
17
20
21
23
24
26
27
32
34
36
37
39
40
41
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TABLES AND FIGURE
5-1
5-2
6-1
6-2
6-3
7-1
7-2
7-3
8-1
9-1
9-2
9-3
9-4
9-5
9-6
9-7
9-8
9-9
9-10
10-1
xii
Nutrient Requirements of White Pekin Ducks as Percentages or Units per Kilogram of Diet
Approximate Body Weights and Feed Consumption of White Pekin Ducks to 8 Weeks of Age
Nutrient Requirements of Ring-Necked Pheasants as Percentages or Units per Kilogram of Diet
Nutrient Requirements of Japanese Quail (Coturnix) as Percentages or Units per Kilogram of Diet
Nutrient Requirements of Bobwhite Quail as Percentages or Units per Kilogram of Diet
Biochemical and Physiological Measurements for Diagnosis of Nutrient Deficiencies in Chickens and Turkeys
Signs of Deficiency in the Embryo
Nutrients Associated with Various Signs of Deficiency in Growing Birds
Toxic Dietary Concentrations of Inorganic Elements and Compounds for Poultry
Composition (Excluding Amino Acids) of Some Feeds Commonly Used for Poultry (data on as-fed basis)
Amino Acid Composition of Some Feeds Commonly Used for Poultry (data on as-fed basis)
Ranges in Weights per Unit of Volume for Selected Feedstuffs at Standard Moisture
Estimation of Amino Acids from Protein Content of Feed Ingredients
Estimation of Amino Acid Composition of Feed Ingredients from Proximate Components
True Digestibility Coefficients (percent) for Selected Amino Acids in Poultry Feedstuffs
Nitrogen Concentration, Crude Protein Equivalents, and Nitrogen-Corrected Metabolizable Energy Values
for Amino Acids
Average Fatty Acid Composition of Some Feeds Commonly Used for Poultry (data on as-fed basis)
Characteristics and Metabolizable Energy of Various Sources of Fats and Selected Carbohydrates Occurring
in Feed
Element Concentrations in Common Mineral Sources (data on as-fed basis)
Formulas for Reference Diets for Chicks
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42
43
44
45
45
47
48
49
59
62
66
68
71
72
74
75
75
76
78
81
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TABLES AND FIGURE
xiii
Figure
1-1
Disposition of dietary energy ingested by a laying hen
4
Appendix Tables
A-1
A-2
A-3
A-4
A-5
A-6
A-7
A-8
A-9
A-10
A-11
A-12
B-1
C-1
Documentation of Nutrient Requirements of Starting and Growing Leghorn-Type Chickens
Documentation of Nutrient Requirements of Leghorn-Type Chickens in Egg Production
Documentation of Nutrient Requirements of Starting and Growing Market Broilers
Documentation of Nutrient Requirements of Broiler Breeder Pullets and Hens
Documentation of Nutrient Requirements of Broiler Breeder Males
Documentation of Nutrient Requirements of Turkeys
Documentation of Nutrient Requirements of Turkey Breeders
Documentation of Nutrient Requirements of Geese
Documentation of Nutrient Requirements of Ducks
Documentation of Nutrient Requirements of Pheasants
Documentation of Nutrient Requirements of Japanese Quail
Documentation of Nutrient Requirements of Bobwhite Quail
Estimating the Energy Value of Feed Ingredients from Proximate Composition
Conversion Factors—Weights and Measures
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85
88
90
97
98
99
105
106
107
109
110
112
113
114
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xiv
Nutrient Requirements of Poultry
Ninth Revised Edition, 1994
Copyright © National Academy of Sciences. All rights reserved.
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xv
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OVERVIEW
1
Overview
The ninth revised edition of Nutrient Requirements of Poultry contains substantially more information than previous
editions. In addition to presenting updated nutrient requirements data, this edition includes more discussion on key facets of
nutrients, nutrient requirements, and nutrient sources. Detailed documentation of the scientific literature used to establish or
estimate the requirements is also included in Appendix A.
Scientifically based knowledge about many nutrient requirements is incomplete. Consequently, calculations and
interpolations were necessary to derive estimated requirements. These nutrient requirements were derived mostly from
empirical observations of responses of poultry to changes in dietary concentrations or intakes of specific nutrients. In some
instances, nutritional models were used to estimate amino acid requirements.
Few nutritional models are available for poultry, primarily because data to support the development of these models are
scarce. There are, however, modeling equations for estimating the energy and amino acid requirements of poultry. Hurwitz et
al. (1978) integrated the energy and amino acid needs of broiler chicks to develop a mathematical model for predicting amino
acid requirements. Models for estimating the amino acid requirements of growing turkeys were proposed by Fisher (1982a)
and Hurwitz et al. (1983a). Modeling equations also have been developed for predicting the energy requirements (National
Research Council, 1987a) and amino acid requirements (Hurwitz and Bornstein, 1973) of laying hens. Additional research is
needed to determine maintenance requirements and partial efficiency of nutrient use for growth versus egg production.
Energy, specific nutrients, and certain nonnutritive feed ingredients are discussed in general terms in Chapter 1.
Definitions of terms used to describe the energy value of poultry feeds are given, and an expanded section on procedures for
determining and estimating dietary metabolizable energy is provided. General aspects of protein and amino acid nutrition and
metabolism have been updated. The section on fats includes information on sources, factors affecting metabolizable energy
(MEn) values, effects on composition of poultry products, and metabolic functions. Overviews are given for minerals,
vitamins, and water. Data on water consumption for chickens and turkeys have been revised according to recent field
observations of contemporary breeds and strains. General characteristics and uses of xanthophylls, unidentified growth
factors, and antimicrobials in poultry diets also are discussed.
Nutrient requirements for specific types of poultry are presented and discussed in Chapters 2 through 6, with each
chapter devoted to a different type. Each of these chapters contains a table or tables detailing the nutrient requirements of the
respective groups. Requirements data are presented on the basis of 90 percent dietary dry matter, which approximates most
feeding conditions. These data are also presented on the basis of total concentrations in the diet or total consumed per day,
not on an available or digestible basis.
In the tables, requirements that are well delineated in the literature, the ''established requirement," are set in regular type.
"Estimated requirements," made on the basis of meager data or by interpolation, are set in bold italicized type. In some
instances, the committee decided to insert a question mark rather than make estimates with no bases.
The committee emphasizes that the requirements values reported herein have not been increased by a "margin of safety."
The values represent the judgment of the subcommittee after its review of the published data. Criteria of adequacy included
growth, reproduction, feed efficiency, health, and quality of poultry products.
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OVERVIEW
2
Ambient temperature and other environmental factors usually were not specified in papers presenting requirements data.
Most experiments, however, have been conducted under moderate conditions, with temperatures of 16° to 21°C and relative
humidities of 40 to 60 percent. When temperature or humidity conditions deviate from these ranges, adjustments in nutrient
concentrations may be needed to compensate for changes in feed intake.
Chapter 2, on the nutrient requirements of chickens, has been divided according to Leghorn-type and meat-type fowl.
For the former, sections are included for starting and growing pullets and for hens in egg production. Similarly, for the latter,
separate sections are presented for starting and growing market broilers, broiler breeder pullets and hens, and broiler breeder
males. Requirements of starting and growing turkeys and turkey breeders are given in Chapter 3. Nutrient requirements of
geese, ducks, and pheasants and quail are provided in Chapters 4, 5, and 6, respectively. These data, however, were based on
a relatively meager amount of literature.
Chapter 7, on signs of nutritional deficiencies in chickens and turkeys, has been enlarged considerably to include more
descriptive information and documentation. Tables present biochemical and physiological indicators of nutrient deficiencies,
signs of nutrient deficiencies in embryos, and nutrient deficiencies that may be associated with specific deficiency signs.
Chapter 8 includes an update presentation on toxic levels of elements as related to diets or drinking water.
Feedstuff composition data and related information are presented in Chapter 9. The tabular data of Tables 9-2 and 9-3
have been revised according to recent analytical results obtained with contemporary feedstuffs. This revision primarily
involved changes in proximate and amino acid compositions of numerous feedstuffs. True metabolizable energy (TMEn)
values of many feedstuffs also have been included in Table 9-2. Two new sections have been added to Chapter 9. One section
briefly discusses and presents equations estimating amino acid composition on the basis of protein content or proximate
analysis. The second covers amino acid availability and includes a listing of true digestibility coefficients for selected amino
acids in many poultry feedstuffs. The tabular presentation in Chapter 9 on fatty acid composition and MEn values of dietary
fats for poultry is extensive and well documented. Information on the crude protein equivalents and nitrogen-corrected MEn
values of amino acids and on the element concentrations in common mineral sources also is provided.
The nutrient composition of feedstuffs is, of course, variable. In addition, the effective concentrations of nutrients in
diets may be reduced by inadequate feed mixing, improper processing, and unfavorable storage conditions. Nutritionists may
accordingly add a "margin of safety" to the stated requirements in arriving at nutrient allowances to be used in formulation to
compensate for these aforementioned conditions.
Examples of practical, semipurified, and chemically defined reference diets for chicks are given in Chapter 10.
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1
Components of Poultry Diets
Poultry diets are composed primarily of a mixture of several feedstuffs such as cereal grains, soybean meal, animal byproduct meals, fats, and vitamin and mineral premixes. These feedstuffs, together with water, provide the energy and
nutrients that are essential for the bird's growth, reproduction, and health, namely proteins and amino acids, carbohydrates,
fats, minerals, and vitamins. The energy necessary for maintaining the bird's general metabolism and for producing meat and
eggs is provided by the energy-yielding dietary components, primarily carbohydrates and fats, but also protein.
Poultry diets also can include certain constituents not classified as nutrients, such as xanthophylls (that pigment and
impart desired color to poultry products), the "unidentified growth factors" claimed to be in some natural ingredients, and
antimicrobial agents (benefits of which may include improvement of growth and efficiency of feed utilization). Each of these
components of poultry diets is considered in the following sections.
ENERGY
Energy is not a nutrient but a property of energy-yielding nutrients when they are oxidized during metabolism. The
energy value of a feed ingredient or of a diet can be expressed in several ways. Thus, a description is presented below of
terminology associated with dietary energy values, including units of measure (digestible energy, metabolizable energy, etc.).
Because metabolizable energy values are most commonly used to define the dietary energy available to poultry, several
procedures for determining metabolizable energy values, by using bioassays or estimates based on proximate analysis, are
described. An example of the disposition of dietary energy ingested by a laying hen and some general considerations
regarding setting dietary energy concentrations of diets follow. Finally, some caveats are given concerning the energy values
listed in the nutrient requirement tables in this report.
Energy Terminology
Energy terms for feedstuffs are defined and discussed in detail in Nutritional Energetics of Domestic Animals and
Glossary of Energy Terms (National Research Council, 1981b). For a more in-depth discussion of energy terms related
specifically to poultry, the reader is referred to Pesti and Edwards (1983). A brief description of the terms most frequently
used in connection with poultry feeds appears below.
A calorie (cal) is the heat required to raise the temperature of 1 g of water from 16.5° to 17.5° C. Because the specific
heat of water changes with temperature, however, 1 cal is defined more precisely as 4.184 joules.
A kilocalorie (kcal) equals 1,000 cal and is a common unit of energy used by the poultry feed industry.
A megacalorie (Mcal) equals, 1,000,000 cal and is commonly used as a basis for expressing requirements of other
nutrients in relation to dietary energy.
A joule (J) equals 107 ergs (1 erg is the amount of energy expended to accelerate a mass of 1 g by 1 cm/s). The joule has
been selected by Le Systéme International d'Unites (SI; International System of Units) and the U.S. National Bureau of
Standards (1986) as the preferred unit for expressing all forms of energy. Although the joule is defined in mechanical terms
(that is, as the force needed to accelerate a mass), it can be converted to calories. The joule has replaced the calorie as the unit
for energy in nutritional work in many countries and in most scientific journals. In this publication, however, calorie is used
because it is the standard energy
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terminology used in the U.S. poultry industry and there is no difference in accuracy between the two terms.
A kilojoule (kJ) equals 1,000 J.
A megajoule (MJ) equals 1,000,000 J.
Gross energy (E) is the energy released as heat when a substance is completely oxidized to carbon dioxide and water.
Gross energy is also referred to as the heat of combustion. It is generally measured using 25 to 30 atmospheres of oxygen in a
bomb calorimeter.
Apparent digestible energy (DE) is the gross energy of the feed consumed minus the gross energy of the feces. (DE =
[E of food per unit dry weight × dry weight of food] - [E of feces per unit dry weight × dry weight of feces]). Birds excrete
feces and urine together via a cloaca, and it is difficult to separate the feces and measure digestibility. As a consequence, DE
values are not generally employed in poultry feed formulation.
Apparent metabolizable energy (ME) is the gross energy of the feed consumed minus the gross energy contained in
the feces, urine, and gaseous products of digestion. For poultry the gaseous products are usually negligible, so ME represents
the gross energy of the feed minus the gross energy of the excreta. A correction for nitrogen retained in the body is usually
applied to yield a nitrogen-corrected ME (MEn) value. MEn, as determined using the method described by Anderson et al.
(1958), or slight modifications thereof, is the most common measure of available energy used in formulation of poultry feeds.
True metabolizable energy (TME) for poultry is the gross energy of the feed consumed minus the gross energy of the
excreta of feed origin. A correction for nitrogen retention may be applied to give a TMEn value. Most MEn values in the
literature have been determined by assays in which the test material is substituted for part of the test diet or for some
ingredient of known ME value. When birds in these assays are allowed to consume feed on an ad libitum basis, the MEn
values obtained approximate TMEn values for most feedstuffs.
Net energy (NE) is metabolizable energy minus the energy lost as the heat increment. NE may include the energy used
for maintenance only (NEm) or for maintenance and production (NEm+p). Because NE is used at different levels of efficiency
for maintenance or the various productive functions, there is no absolute NE value for each feedstuff. For this reason,
productive energy, once a popular measure of the energy available to poultry from feedstuffs and an estimate of NE, is
seldom used.
Disposition of Dietary Energy
Figure 1-1 illustrates the proportional relationships in the disposition of dietary energy ingested by a laying hen. Energy
is voided or used at various stages following consumption of 1 kg feed by the hen.
Figure 1-1 Disposition of dietary energy ingested by a laying hen.
Of 4,000 kcal provided in 1 kg of this particular diet, 2,900 kcal are capable of being metabolized by the hen and about
2,300 kcal are available for maintenance and transfer into body tissue and egg (net energy) (Fraps, 1946; Hill and Anderson,
1958; Titus, 1961). The relative amounts of both metabolizable and net energy will, of course, vary with the composition of
the feedstuffs in the diet. Other factors, such as the species, genetic makeup, and age of poultry, as well as the environmental
conditions, also influence the precise distribution of dietary energy into the various compartments (Scott et al., 1982).
Procedures for Determining Metabolizable Energy
Metabolizable energy is determined by various bioassay procedures whereby feed intake and excreta output are related
over a 2- to 5-day test period. Apparent metabolizable energy is most commonly determined through actual measurement of
feed intake and excreta output, or by determining the ratio of dry matter intake to output through use of an inert dietary
marker, such as chromic oxide (Cr2O3). A number of potential problems arise with use of markers (Kane et al., 1950; Vohra
and Kratzer, 1967; Duke et al., 1968; Vohra, 1972a), and thus the latter method often leads to more variation in final
determined ME values (Potter, 1972).
When the ME value of an ingredient is to be determined, two or more diets must be used, since feeding an ingredient by
itself can cause palatability problems and fails to accommodate potential synergism between nutrients. The two methods most
frequently used in substituting the test ingredient into a control basal diet are those described by Anderson et al. (1958) and
Sibbald and Slinger (1963). In the former method the test ingredient is substituted for glucose, but in the latter method the test
ingredient is substituted for all the energy-yielding ingredients of the basal diet. Anderson et al. (1958) proposed that the
value of 3.65 kcal/g be
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used as the standard for glucose. The basal diet used by Anderson et al. (1958), containing about 50 percent glucose and
designated as E9, has been used extensively in determinations of nitrogen-corrected ME (MEn).
In the method of Sibbald and Slinger (1963) the test ingredient is substituted essentially for part of the complete basal
diet. However, to avoid mineral and vitamin deficiencies, components of the diet containing these nutrients are left intact,
The use of two basal diets of differing protein contents was proposed to maintain the protein contents of substituted diets
within an acceptable range. An advantage of the substitution method of Sibbald and Slinger (1963) is that the MEn value of
the reference basal diet is necessarily determined in each MEn assay. Although samples of glucose are likely to be less
variable than samples of regular feed ingredients, the MEn of glucose may vary under different dietary conditions, and its
MEn value should be determined under the experimental conditions used (Mateos and Sell, 1980).
The test ingredient may be substituted at one or more levels. Regardless of the basal diet used, the accuracy of the MEn
value obtained depends to some extent on the proportions of the test ingredient substituted into test diets. In extrapolating to
calculate the MEn value of the test ingredient, the error of determination of the test ingredient is therefore multiplied by a
factor of 100 divided by percentage of substitution. Therefore the highest proportion of the test ingredient possible in the test
diet should be used. Usually, this amount is determined by nutrient balance and palatability.
Potter et al. (1960) proposed a linear regression procedure for the calculation of MEn values for ingredients substituted at
several levels. The ingredient MEn value is derived by extrapolation to 100 percent inclusion from a regression equation
relating test diet MEn values and proportion of test ingredient in such diets. As for most other methods of MEn determination,
a criticism of the regression methods is that the extrapolation is beyond the range of experimental data. Sibbald and Slinger
(1962) pointed out that this general criticism is of little significance as long as the range of inclusion levels used is within that
normally encountered under practical conditions because it is the application of ingredient MEn values in commercial dietary
formulation that is of interest.
TME was described as an estimate of ME in which correction is made for metabolic fecal and endogenous urinary
energy (National Research Council, 1981b). These energy components of excreta are not directly of dietary origin, and, as
suggested by Sibbald (1980), correction for their excretion in bioassays leads to TME. It should be noted that ME as
determined using the procedure of Anderson et al. (1958) inherently corrects for metabolic fecal and endogenous urinary
energy excretion, whereas the method of Sibbald (1976) for determining ME does not. The TME method is quite rapid in that
it takes only a 48-hour collection period and, because ingredients are force-fed, there is no need to use a series of basal and
test diets.
The TME procedure, however, has been subjected to criticism. TME determinations assume that fecal metabolic and
urinary endogenous energy excretions are constant, irrespective of feed intake. Data have been presented showing that, to the
contrary, metabolic and endogenous energy excretions are influenced by amount and nature of materials passing through the
gastrointestinal tract (Farrell, 1981; Farrell et al., 1991; Tenesaca and Sell, 1981; Hartel, 1986). Another criticism is that
ingredients are often force-fed alone, thereby preventing synergistic or antagonistic effects between or among ingredients on
energy utilization. Synergism is known to occur between fatty acids (Young, 1961; Artman, 1964; Leeson and Summers,
1976a) and there is evidence for synergism between protein concentrates (Woodham and Deans, 1977). A third criticism of
the TME method relates to the imposition of 48 periods of feed deprivation, which would result in an abnormal physiological
status of the bird.
Both ME and TME should be corrected for nitrogen retention that occurs during the assay period. If, during an ME
determination, nitrogen is retained by the animal, the excreta will contain less urinary nitrogen and hence less energy would
be excreted as compared with an animal that is not retaining N. Because the extent of nitrogen retention differs with age and
species, a correction factor is essential if comparisons of ME values for the same ingredient with different animals are to be
made.
Hill and Anderson (1958), assuming that if nitrogen is not retained it will appear as uric acid, proposed a correction
value of 8.22 kcal/g nitrogen retained because this is the energy obtained when uric acid is completely oxidized. This
assumption has been criticized because only 60 to 80 percent of the nitrogen of chicken urine is in the form of uric acid
(Coulson and Hughes, 1930). However, the assumption that oxidation of varying amounts of protein would yield a consistent
pattern of nitrogenous excretory products seems no more correct than the assumption that all nitrogen would be excreted as
uric acid (Hill and Anderson, 1958). Thus, from a practical viewpoint, the uric acid value has been used most frequently and
is generally quoted (Scott et al., 1982).
Sibbald and Slinger (1963) questioned the validity of correcting for nitrogen retention, suggesting that correction does
little to improve the usefulness of classical ME values and that the extra work involved is not justified. Potter (1972),
however, suggested that correction to zero nitrogen retention is essential for reproducible results when the MEn of a single
diet is to be measured
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with birds of various ages because of differences in rates of protein accretion or protein catabolism. Correction to a speciesspecific or age-specific nitrogen retention, although having the advantage of applicability for specific circumstances, cannot
be used in comparative work because "typical" nitrogen retention varies with species and age. Leeson et al. (1977a) indicated
the need for nitrogen correction in interpretation of bioassay data.
An alternative to classical bioassay is based on changes in rate of growth in response to dietary energy. Squibb (1971)
suggested a method for the "standardization and simplification" of MEn determination procedures. The method is a
modification of that described by Yoshida and Morimoto (1970). It is based on the premise that rapidly growing immature
animals restricted in terms of energy intake but given adequate protein will show an increase in growth in direct proportion to
energy added to the diet. Considering the restricted feeding of the energy-deficient diet used by Squibb (1971), the adequacy
of the protein in terms of quantity and quality can be questioned. However, the concept warrants further study as a means of
evaluating the energy value of ingredients, such as fats, that are difficult to assay using conventional procedures.
Most MEn values reported for feedstuffs have been determined with young chicks. Although adult male chickens have
been used to determine TMEn content of many feedstuffs, few studies have been done to determine either MEn or TMEn for
poultry of different ages. More MEn and TME n data are needed for many feed ingredients for chickens, turkeys, and other
poultry of different ages.
Estimation from Proximate Composition
Several researchers have developed prediction equations to estimate the energy content of feed ingredients from their
proximate components. Prediction of the "usable" energy value of a feed from its chemical composition has been attempted
for many years. The Weende, or proximate analysis, system was developed as an attempt to predict the nutritional value
(including the energy value) of an ingredient or of mixed feed from its component parts. Fraps et al. (1940) predicted the ME
content of feeds from the values for digestible crude protein, ether extract, and nitrogen-free extract (NFE). Titus (1955) used
this concept to derive a series of "percentage multipliers" for the calculation of ME values for different types of feed
ingredients. Later, these ''percentage multipliers" were updated and extended to a wider range of ingredients (Titus and Fritz,
1971).
Janssen et al. (1979) conducted a series of studies to correlate the chemical composition of different types of feed
ingredients to the ME value. By using multiple regression analysis, equations were derived to estimate MEn (kcal/kg dry
matter) from chemical composition. More recently, a subcommittee of the European Federation of the World's Poultry
Science Association (1989) developed a set of equations to estimate the energy value of ingredients. Data sets from a number
of European laboratories were combined to develop the equations. A list of prediction equations that have been published
recently is provided in Appendix Table B-1. Dale et al. (1990) developed an equation to estimate the TMEn value of dried
bakery products, a blend of various by-products produced by the baking industry.
The ME value of grain sorghums is known to be influenced by their tannin content. Sibbald (1977) reported TME values
of 3,300 and 3,970 kcal/kg for high- and low-tannin grain sorghums, respectively, and Queiroz et al. (1978) found MEn
values of 2,886 and 3,091 kcal/kg for high- and low-tannin grain sorghums. Gous et al. (1982) found a highly significant
negative correlation between the MEn of grain sorghums and their tannic acid content, the relationship due to a decreased
digestibility with increasing tannic acid concentration. These researchers developed a regression equation to estimate ME
from tannic acid concentration. A similar equation was developed by the European Federation of the World's Poultry Science
Association in 1989. Although these equations may result in slightly different estimates, they both point out the adverse
effects of the tannin content on digestibility of grain sorghums.
Moir and Connor (1977) developed equations to predict MEn of grain sorghums using three different types of crude fiber
assays. The ME n content of sorghum was predicted from the three fiber assay methods with precision of, respectively, ±117,
±148, and ±126 kcal/kg dry matter. These values correspond to coefficients of variation of 3.0, 3.8, and 3.3 percent,
respectively. Thus, any of the three fiber methods could be used to predict the MEn of grain sorghums for poultry.
Considerable variation exists in the nutrient composition of poultry by-product meal from various production lots and
among producers, depending on raw material used (e.g., proportions of feet, legs, blood, and offal may vary considerably).
Pesti et al. (1986) determined the TMEn of a number of samples of poultry by-product and derived several equations to
estimate TMEn from various measurements. The equations vary in complexity, some using only one parameter to estimate
TMEn and others using two measurements. The coefficients of determination (R2) for the two-measurement equations were
similar; thus, persons using these equations may select measurements that are in concert with the capability of their own
laboratory.
Perhaps the most difficult feed ingredients to analyze for MEn are supplemental fats. Many factors influence the
digestibility and subsequent MEn of fats; these have
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been extensively reviewed by Renner and Hill (1961), Young and Garrett (1963), Lewis and Payne (1966), Hakansson
(1974), Leeson and Summers (1976a), Fuller and Dale (1982), Ketels et al. (1987), Ketels and DeGroote (1988), and many
others. Prominent among these factors are age of poultry, level of fat inclusion in the diet, and overall fatty acid composition
of the diet. Several studies have been conducted to estimate the energy value of a fat from its composition. Janssen et al.
(1979) estimated the energy value of fats produced by Dutch renderers (Appendix Table B-1). Huyghebaert et al. (1988)
evaluated a wide variety of fats and developed prediction equations for MEn using multiple linear regression analysis
involving different characteristics of fats. Several equations were developed for (1) all fats and oils examined and (2)
different categories of fats (e.g., animal or vegetable fats). The accuracy of the equations was improved by separating the fats
into different categories.
It is well known that utilization of saturated fatty acids is improved by the presence of unsaturated fatty acids in the fat
blend (Young and Garrett, 1963; Young, 1965; Lewis and Payne, 1966; Garrett and Young, 1975; Leeson and Summers,
1976a). The nature of the fat in the basal diet has a significant effect on the utilization of supplemental fats (Sell et al., 1976;
Sibbald and Kramer, 1978; Fuller and Dale, 1982). These interactions between the supplemental fat and the basal dietary fat
are especially noticeable at low inclusion levels of supplemental fat (Wiseman et al., 1986; Ketels et al., 1987).
Ketels and DeGroote (1989) evaluated the relationship between the ratio of unsaturated to saturated fatty acids (U:S) in
the diet and MEn of a number of fats and developed equations relating fat MEn, fat utilization, and the utilization of specific
fatty acids to the U:S for young broiler chickens. Best fit regression equations for supplemental fat utilization and fat MEn
were exponential. Fat utilization increased rapidly in the U:S range of 0 to 2.5, reaching a near-asymptotical maximum at a
U:S of 4. Synergism between added fats, due either to blending vegetable oils with animal fats or to using basal diets with
unsaturated lipid fractions, led to increased utilization of animal fats. Utilization of vegetable oils was not influenced by
changing U:S ratios. The effect of factors influencing fat utilization, such as level of supplemental fat and basal diet
composition, seemed to be primarily through variation in degree of saturation of the total dietary lipid fraction. For young
broilers, about 75 percent of the variation in fat utilization and MEn was due to differences in the chemical composition of the
fat fraction.
Excellent summaries of the use of indirect methods for estimating the ME in feed ingredients have been presented by
Harris et al. (1972), Sibbald (1975, 1982), Eackhout and Moermans (1981), Fisher (1982b), Fonnesbeck et al. (1984), and
Just et al. (1984). These reports discuss many of the problems associated with the use of indirect procedures to replace
conventional bioassays for ME.
At this time, the committee cannot recommend the best equation(s) to use to estimate ME from chemical composition.
To date, no studies have compared the various equations with a determined value. In addition, some of the chemical
determinations are subject to much variability or are relatively complex and may not be easy to adapt to some laboratory
situations. Users may wish to calculate ME by using as many of the equations as seem feasible and then evaluating the results
before selecting the procedure that is most appropriate for their situation.
Setting Dietary Levels
In formulating poultry diets, energy level is usually selected as the starting point. An appropriate energy level is one that
most likely results in the lowest feed cost per unit of product (weight gain or eggs). The feed cost per unit of product, in turn,
is determined by the cost per unit weight of diet and the amount of diet required to produce a unit of product. In areas of the
world where high-energy grains and feed-grade fats are relatively inexpensive, high-energy diets are often most economical
(i.e., the lowest feed cost per unit of product); however, if a leaner carcass is desired, it may be necessary to consider other
levels of dietary energy. In areas where lower-energy grains and by-products are less expensive, low-energy diets are often
most economical.
The dietary energy level selected is often used as a basis for setting most nutrient concentrations in a diet. This approach
to formulation of poultry diets is based on the concept that poultry tend to eat to meet their energy needs, assuming that the
diet is adequate in essential nutrients (Hill and Dansky, 1950; 1954; Hill et al., 1956; Scott et al., 1982). Such an assumption,
however, must be used with caution and with an understanding of its potential limitations. For example, if a diet is deficient
in any nutrient, daily feed consumption may decrease in relation to the severity of the deficiency. One exception may occur
with an amino acid deficiency, whereby a marginal deficiency may result in a small increase in feed consumption. If a diet
has a gross excess of any nutrient, daily feed consumption usually decreases in relation to the severity of the potential toxicity.
The physiological mechanisms by which poultry respond to different dietary energy concentrations are not known,
although several possible mechanisms have been proposed (National Research Council, 1987a). Equations that can be used to
predict feed and energy
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intakes of laying hens and coefficients to predict the energy requirements of broiler chickens have been given by the National
Research Council (1987a).
Although poultry generally adjust feed consumption to achieve a minimum energy intake from diets containing different
energy levels, these adjustments are not always precise. Morris (1968) summarized data from 34 experiments and found that
laying hens overconsumed energy when fed high-energy diets, and the degree of overconsumption was greatest for strains
with characteristically high-energy intakes. Data from a large number of broiler chicken experiments also showed that
changes in feed intake were not inversely proportional to changes in dietary energy level, especially when broilers were fed
moderateto high-energy diets (Fisher and Wilson, 1974). More recent studies also illustrated that growing broilers and
turkeys consume more energy when fed high-energy diets than those fed low- to moderate-energy diets (Sell et al., 1981;
Owings and Sell, 1982; Sell and Owings, 1984; Brue and Latshaw, 1985; Potter and McCarthy, 1985). For laying hens, some
combinations of carbohydrates, fat, and protein resulted in more energy intake than others (Rising et al., 1989). Diets with 3
percent fat increased daily feed intake in comparison with diets containing no added fat, and hens fed diets that provided
more protein also consumed greater amounts of energy. Generally, regulation of energy intake by laying hens and broilers is
more precise when relatively low-energy diets are fed (Morris, 1968; Fisher and Wilson, 1974; Latshaw et al., 1990). In some
instances, however, laying hens are fairly accurate in regulating energy consumption when fed high-energy diets (Horani and
Sell, 1977).
Because the preponderance of data shows that changes in feed intake usually are not proportional to changes in dietary
energy concentration, the use of specific protein/amino acid-to-dietary energy ratios (originally termed energy-to-protein
ratios) in formulating poultry diets (Baldini and Rosenberg, 1955; Combs, 1961; Scott et al., 1982; Thomas et al., 1986) must
be carefully evaluated. Relating nutrient concentrations to dietary energy level seems to have greatest practical application for
Leghorn chickens that generally are fed diets of low to moderate energy content. In the instance of growing broiler chickens
and turkeys, however, maintaining specific nutrient-to-energy ratios seems questionable. This is particularly true for proteinto-energy ratios intended to support economical growth and feed efficiency (Pesti and Fletcher, 1983; Sell et al., 1985; 1989).
If the production of lean broiler or turkey carcasses is of economic importance, appropriate dietary protein-to-energy ratios
may be of greater significance. It would be desirable to have mathematical models available that would facilitate the selection
of most economical combinations of dietary concentrations of protein/amino acids (and other nutrients) and energy to achieve
poultry production goals. Development of such models will be contingent on research designed to obtain more relevant
information than is currently available.
Factors other than dietary energy and nutrient balance that affect feed intake include bulk density of the diet (Cherry et
al., 1983) and ambient temperature (National Research Council, 1981a). The latter can have considerable impact on feed
consumption of poultry, especially adult birds, because feed intake decreases as ambient temperature increases. Leghorn-type
hens consume approximately 1.5 g less feed per hen daily for each 1°C increase in ambient temperature over the range of 10°
to 35°C (Davis et al., 1973; Sykes, 1979). At temperatures above 30°C, the decrease in feed consumption may be 2.5 to 4 g
for each 1°C increase (Sykes, 1979; Sell et al., 1983). Similar responses of decreasing feed intake with increasing
temperatures have been reported for turkeys (Parker et al., 1972; Hurwitz et al., 1980).
Energy Values in the Nutrient Requirement Tables
The MEn values heading the lists of nutrient requirements given in Chapters 3 through 6 should not be regarded as
energy requirements. The committee chose these as bases of reference. They represent the dietary energy concentrations
frequently used under practical conditions of feed formulation and poultry management. For those persons preferring to use
TMEn values, the TMEn values of numerous feed ingredients are included in Table 9-1. Generally, MEn values as determined
by the method of Anderson et al. (1958) and TMEn values as determined by Sibbald (1983) are similar for many ingredients.
However, MEn and TMEn values differ substantially for some ingredients, such as feather meal, rice bran, wheat middlings,
and corn distillers' grains with solubles, and so in these instances MEn values should not be indiscriminately interchanged
with TMEn values for purposes of diet formulation.
CARBOHYDRATES
Dietary carbohydrates are important sources of energy for poultry. Cereal grains such as corn, grain sorghum, wheat,
and barley contribute most of the carbohydrates to poultry diets. The majority of the carbohydrates of cereal grains occurs as
starch, which is readily digested by poultry (Moran, 1985a). Other carbohydrates occur in varying concentrations in cereal
grains and protein supplements. These carbohydrates include polysaccharides, such as cellulose, hemicellulose, pentosans,
and oligosaccharides, such as stachyose and raffinose, all of which are poorly digested by poultry. Thus, these dietary
carbohydrates often
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contribute little to meeting the energy requirement of poultry, and some adversely affect the digestive processes of poultry
when present in sufficient dietary concentrations. For example, the pentosans of rye and beta glucans of barley increase the
viscosity of digesta and thereby interfere with nutrient utilization by poultry (Wagner and Thomas, 1978; Antoniou and
Marquardt, 1981; Classen et al., 1985; Bedford et al., 1991). Supplementation of rye or barley-containing diets with
appropriate supplemental enzyme preparations improves nutrient utilization and growth of young poultry (Leong et al., 1962;
Edney et al., 1989; Friesen et al., 1992).
PROTEINS AND AMINO ACIDS
Dietary requirements for protein are actually requirements for the amino acids contained in the dietary protein. Amino
acids obtained from dietary protein are used by poultry to fulfill a diversity of functions. For example, amino acids, as
proteins, are primary constituents of structural and protective tissues, such as skin, feathers, bone matrix, and ligaments, as
well as of the soft tissues, including organs and muscles. Also, amino acids and small peptides resulting from digestionabsorption may serve a variety of metabolic functions and as precursors of many important nonprotein body constituents.
Because body proteins are in a dynamic state, with synthesis and degradation occurring continuously, an adequate intake of
dietary amino acids is required. If dietary protein (amino acids) is inadequate, there is a reduction or cessation of growth or
productivity and a withdrawal of protein from less vital body tissues to maintain the functions of more vital tissues.
There are 22 amino acids in body proteins, and all are physiologically essential. Nutritionally, these amino acids can be
divided into two categories: those that poultry cannot synthesize at all or rapidly enough to meet metabolic requirements
(essential) and those than can be synthesized from other amino acids (nonessential). The essential amino acids must be
supplied by the diet. If the nonessential amino acids are not supplied by the diet, they must be synthesized by poultry. The
presence of adequate amounts of nonessential amino acids in the diet reduces the necessity of synthesizing them from
essential amino acids. Thus, stating dietary requirements for both protein and essential amino acids is an appropriate way to
ensure that all amino acids needed physiologically are provided.
Variations in Requirements
Protein and amino acid requirements vary considerably according to the productive state of the bird, that is, the rate of
growth or egg production. For example, turkey poults and broiler chickens have high amino acid requirements to meet the
needs for rapid growth. The mature rooster has lower amino acid requirements than does the laying hen, even though its body
size is greater and its feed consumption is similar.
Body size, growth rate, and egg production of poultry are determined by their genetics. Amino acid requirements,
therefore, also differ among types, breeds, and strains of poultry, as can be seen by comparing the values shown in the
requirement tables provided in this report for the different types of poultry. Genetic differences in amino acid requirements
may occur because of differences in efficiency of digestion, nutrient absorption, and metabolism of absorbed nutrients
(National Research Council, 1975).
Although dietary requirements for amino acids and protein usually are stated as percentages of the diet, the quantitative
needs of poultry must be met by a balanced source to obtain maximum productivity. Thus factors that affect feed
consumption also will affect quantitative intakes of amino acids and protein, and, consequently, will influence the dietary
concentration of these nutrients needed to provide adequate nutrition. Factors affecting feed consumption are discussed in the
section on "Setting Dietary Levels" and have been reviewed in the National Research Council (1987a) publication, Predicting
Feed Intake of Food-Producing Animals.
As discussed in the section "Setting Dietary Levels," adjustments in the protein and amino acids concentration of diets
may be necessary to compensate for difference in energy concentration of diets. This is especially true for White Leghorn
chickens (Morris, 1968; Byerly et al., 1980) and turkey hens (Kratzer et al., 1976).
Ambient temperature also affects feed intake of poultry (Hurwitz et al., 1980). Protein and amino acid requirements
listed herein generally pertain to poultry kept in moderate temperatures (18° to 24°C). Ambient temperatures outside of this
range cause an inverse response in feed consumption; that is, the lower the temperature, the greater the feed intake and vice
versa (National Research Council, 1981c). Consequently, percentage requirements of protein and amino acids should be
increased in warmer environments and decreased in cooler environments, in accordance with expected differences in feed
intake. These adjustments may aid in ensuring required daily intakes of amino acids. Some precautions, however, should be
used in increasing the dietary protein concentration for poultry subjected to high ambient temperature. Waldroup et al.
(1976d) reported that performance of broiler chicks was improved by minimizing excess dietary amino acids.
Information available from research documenting the influence of dietary energy concentration and ambient
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temperature on feed intake has been integrated with data describing amino acid needs for maintenance, body growth (such as
for muscle and feathers), or egg production to derive mathematical models to predict the dietary amino acid requirements of
poultry (Fisher et al., 1973; Hurwitz and Bornstein, 1973; Hurwitz et al., 1978; Emmans, 1981; Slagter and Waldroup, 1984).
Prediction models may be useful in feed formulation, and they also provide valuable insight into areas of amino acid and
protein nutrition where more definitive information is needed on requirements.
Dietary protein concentrations can affect the requirements for individual essential amino acids. Generally, as dietary
protein level increases, essential amino acid requirements (expressed as a percentage of the diet) increase, although when
expressed as a percentage of the protein, essential amino acid requirements are little affected (Almquist, 1952; Boomgaardt
and Baker, 1971, 1973a; Morris et al., 1987; Robbins, 1987; Mendonca and Jensen, 1989a). These observations demonstrate
the importance of maintaining a balance among the concentrations of essential and nonessential amino acids in poultry diets.
Optimal balance is important for efficient utilization of dietary protein.
The protein and amino acid concentrations presented as requirements herein are intended to support maximum growth
and production. Achieving maximum growth and production, however, may not always ensure maximum economic returns,
particularly when prices of protein sources are high. If decreased performance can be tolerated, dietary concentrations of
amino acids may, accordingly, be reduced somewhat to maximize economic returns.
Specific Amino Acid Relationships
Although each amino acid can be metabolized independently of others, relationships between certain amino acids exist.
In some instances, the relationship may be beneficial. For example, one amino acid may be converted to another to fulfill a
metabolic need. In other instances, a metabolic antagonism may exist with undesirable consequences. A brief description of
amino acid relationships that may be of importance in poultry nutrition is given in the following section.
Methionine Plus Cystine
Methionine can donate its methyl group to biological processes, and the resulting sulfur-containing compound,
homocysteine, together with serine, can be used to synthesize cysteine via cystathionine. The sulfhydryl groups of two
molecules of cysteine are oxidized to form cystine. This conversion cannot be reversed, and two methionine molecules are
needed to ultimately supply the two sulfur atoms of cystine (du Vigneaud, 1952; Creek, 1968; Baker, 1976). The requirement
for methionine can be satisfied only by methionine, whereas that for cystine can also be met with methionine.
The catabolism of methionine and cystine largely leads to conversion of the associated sulfur into sulfate. This sulfate
may be used in metabolism, particularly as a part of certain connective tissues. Similarly, methyl groups of methionine may
be used in transmethylation and the de novo synthesis of sarcosine, betaine, and choline. Choline is a constituent of
phospholipids, and its incorporation into membranes is extensive. During rapid growth, when accrual of connective tissue and
expansion of membrane surfaces are great, an increased sensitivity to methionine at levels marginal to the requirement may
occur if dietary choline and sulfate are not sufficient (Baker et al., 1983; Miles et al., 1983; Blair et al., 1986).
Phenylalanine Plus Tyrosine
Tyrosine is the initial product formed during the biological degradation of phenylalanine. In turn, phenylalanine can be
used to meet the bird's need for tyrosine on a mole-for-mole basis (Creek, 1968; Sasse and Baker, 1972). Although this
conversion may be reversed to a small extent and tyrosine used to form phenylalanine, its contribution is too small to be of
practical significance (Ishibashi, 1972).
Glycine Plus Serine
Although glycine can be synthesized by fowl, the rate is not adequate to support maximal growth (Featherston, 1976).
Serine can be converted to glycine on an equimolar basis. This reaction is reversible, and glycine can be used to form serine
(Sugahara and Kandatsu, 1976).
Imbalance, Antagonism, And Toxicity
The essential amino acids are related to one another by virtue of need to support production plus maintenance. The
combined need for production and maintenance represents the bird's requirement. Requirement for any one essential amino
acid represents the combined need for maintenance plus production. Each essential amino acid is unique in its catabolism,
and an inadequacy of any one of them (the first limiting) usually necessitates some catabolism of the others. The bird's
response can vary with the essential amino acid, the extent of its inadequacy, and existing relationships among the remainder.
As an example, Sugahara et al. (1969) fed chicks a purified amino acid diet corresponding to 100 percent of the requirement
for all essential amino acids as the positive control and compared the performance response to when all amino acids were
reduced to 60 percent of the requirement as opposed to 60 percent reduction
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with each one alone. Weight gain was better with individual decreases of methionine-cystine, leucine, lysine, and arginine
than when a total reduction was imposed, whereas additional weight loss occurred with individual decreases of
phenylalanine, tyrosine, tryptophan, isoleucine, valine and threonine. A reduction in dietary histidine gave a similar response
to that observed when all amino acids were reduced.
Deficiencies of any one of the essential amino acids can be exaggerated by adding purified amino acids and/or
combining complete proteins such that the extent of difference between the first and second limiting amino acid increases.
The response is generally an additional impairment of body weight gain. Accentuation of the deficiency in this manner
usually involves diets of low protein content, and a decrease in feed intake is the fundamental reason for poor weight gain
rather than alteration in effectiveness of the first limiting amino acid (Fisher et al., 1960; Fisher and Shapiro, 1961; Netke et
al., 1969).
Amino acid antagonisms may also accentuate a deficiency of the first limiting amino acid, but these differ from
imbalances because utilization of the limiting amino acid is reduced. Antagonisms can occur between amino acids having
side chains exhibiting similar structural and/or chemical characteristics, and increasing the dietary concentration of one that is
in excess of productive use adversely affects metabolism of the other. In a situation in which one essential amino acid is first
limiting, increasing the other's concentration to enlarge the difference antagonizes the use of the first limiting amino acid and
induces or exacerbates a deficiency.
Antagonisms have been shown to exist for leucine-isoleucine-valine, arginine-lysine, and threonine-tryptophan (D'Mello
and Lewis, 1970). The most important of these antagonisms occurs with leucine and isoleucine. Certain feedstuff
combinations (for example, corn plus corn gluten meal) can lead to practical diets in which leucine is at particularly high
levels while isoleucine is marginal in adequacy. Amino acid levels that would be likely to provoke the other antagonisms
probably would not occur in practice unless high levels of supplemental amino acids were used in low-protein diets.
An amino acid toxicity requires a particularly high level of one amino acid relative to all others. Such an occurrence is
unlikely under practical circumstances because differences of sufficient magnitude do not exist in most protein feedstuffs.
Supplemental methionine and lysine are routinely used by the feed industry but usually in quantities low enough to pose no
threat of toxicity.
Errors in amino acid use may lead to toxicities, however. Methionine is toxic when excessive. Ueda et al. (1981)
observed severe depression in feed consumption and growth of chicks given ad libitum access to a diet containing 10 percent
protein and 1.5 percent L-methionine. Force-feeding this high-methionine, low-protein diet in amounts equal to the feed
intake of controls resulted in death of the chicks. Edmonds and Baker (1987) added excesses of several amino acids to a 23
percent protein corn-soybean meal diet for chicks. Methionine at 4 percent of the diet led to a 92 percent reduction in weight
gain, whereas similar excesses of tryptophan, lysine, and threonine were far less toxic.
Amino Acid Conversion to Vitamins
Niacin is the only vitamin that can be synthesized from an amino acid. Tryptophan can be used to alleviate a dietary
niacin deficiency, but the rate of conversion is poor (Baker et al., 1973). When methionine is provided at levels exceeding use
for protein synthesis, the additional methyl groups may decrease the dietary choline requirement (Pesti et al., 1980). Using
amino acids to spare other nutrients is not currently economical under practical conditions.
Amino Acid Availability
It is well known that the availability of amino acids varies greatly among feedstuffs. The importance of considering
amino acid availability in formulation of poultry diets is discussed in Chapter 9.
FATS
Fat is usually added to the feed for meat-type poultry to increase overall energy concentration and, in turn, improve
productivity and feed efficiency. Oxidation of fat is an efficient means to obtain energy for the cell in large quantity, whereas
anabolic use involves direct incorporation into the body as a part of growth. Lipid accrual is most obvious in adipose tissue;
however, cell multiplication also requires an array of lipids to form associated membranes. These two uses can occur
simultaneously; however, the extent of each may vary considerably.
Sources
Feed-grade fat may come from many different sources. Grease from restaurants, the rendering of animal carcasses, and
the refuse from vegetable oil refining are major sources. These sources represent several types and categories, and each is
defined by the Association of American Feed Control Officials (1984). These definitions indicate fat components and limits
of nonfat material (Sell, 1988). Moisture (M) and those
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compounds that are either insoluble in ether (I) or unsaponifiable (U) are usually of no value, and their composite (MIU)
essentially acts as a diluent.
Total fatty acids contributed by all lipid categories, the proportion that are in free form, and the types of fatty acids
present provide information related to expected digestibility as well as how the fat may be used subsequently. Fatty acid
chain length, extent of unsaturation, and nature of esterification all influence intestinal absorption (Moran, 1989a). The
percentage MIU and percentage digestibility combine to influence the MEn value. All feed fats should be stabilized by an
antioxidant to preserve unsaturated fatty acids and routinely monitored for the possible presence of undesirable residues such
as insolubles, chlorinated hydrocarbons, and unsaponifiables and for peroxides (Rouse, 1986).
Metabolizable Energy Value
Factors influencing the MEn value of fat that are not directly associated with fat quality are age of poultry and method of
measurement. Improved utilization of dietary fats has been shown to occur after 2 to 6 weeks of life for chickens (Renner and
Hill, 1960, 1961; Sibbald, 1978a; Lessire et al., 1982) and turkeys (Whitehead and Fisher, 1975; Sell et al., 1986b). This
improvement is particularly evident with long-chain saturated fatty acids and fats containing substantial proportions of these
fatty acids (Young and Garrett, 1963; Sell et al., 1986b).
The methodology used in obtaining feedstuff energy values has an effect on the values obtained. (See the sections above
on procedures for determination of MEn and on estimating the MEn content of ingredients from proximate composition.)
Actual digestibility of fat may also be used to estimate energy content, and Sell et al. (1986b) found that values determined
by this method agree with concurrent MEn measurements.
When the effects of method of determination and age of the bird are superimposed on factors associated with the fat, it
becomes evident that assigning a specific MEn value to a fat may be inappropriate. The information in Table 9-9 provides a
description of fats that may be used in feeds and their MEn values observed under a variety of circumstances. Data indicate
that considerable variation exists and several factors must be considered in determining feeding value. Some of these factors
are included in the equations listed in Appendix Table B-1, which can be used to predict the MEn value of fats.
Blending Fats
When animal tallow is added to feed at a low level, it may be beneficial to blend it with a small amount of vegetable oil.
The resulting MEn value of blends is greater than can be explained from the arithmetic combination. A synergism in the
absorption of the saturated fatty acids related to the added amounts of unsaturated fatty acids is suspected (Ketels et al., 1986;
Ketels and DeGroote, 1987).
The properties of animal tallows also may be enhanced by the presence of feed ingredients that contain unsaturated fatty
acids. Corn is particularly advantageous in this respect because its fatty acids are mostly unsaturated and it usually constitutes
a large portion of a feed. Sibbald and Kramer (1980) noted that the TME for beef tallow was greater when a corn-based
carrier was used during measurement than when wheat was used.
Extra Caloric Effect
Employing high levels of added fat often leads to more MEn than can be accounted for from the summation of
ingredients. High level fat feeding evidently increases the intestinal retention time of feed and so allows for more complete
digestion and absorption of the nonlipid constituents (Mateos and Sell, 1981; Mateos et al., 1982; Sell et al., 1983).
Improved Net Energy of Production
All body tissues have an energy value that corresponds to their heat of combustion. The net energy of production
corresponds to this energy gained from either body growth or egg formation. Adding fat to feed as an isoenergetic
substitution for carbohydrate usually results in an improved productive energy when the same level of MEn has been derived.
Such improvement is particularly obvious through that period preceding adolescent development. Sell and Owings (1984)
noted that added fat increased the body weight gain of large turkeys, with the greatest advantage occurring between 12 and 20
weeks of age. After 20 weeks, the favorable effect of fat on body weight progressively dissipates, but the effect on feed
efficiency remains (Moran, 1982).
Fatty acid synthesis within fowl occurs primarily in the liver. Immediately preceding sexual maturity the rate of
synthesis increases dramatically, and the rate at which the body's depots accrue fat is great (Moran, 1985b). The provision of
fat in feed obviates the cost of synthesis and is more energy-efficient than is synthesis of fat from carbohydrate.
Laying hens also may respond to added dietary fat. Most lipid in egg yolk is formed in the liver by using fatty acids
obtained from the diet or from de novo synthesis. Providing dietary fat decreases the need for hepatic fatty acid synthesis and
generally increases yolk formation and the weight of the egg (Whitehead, 1981;
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March and MacMillan, 1990). Such advantages are particularly valuable during high environmental temperatures. As feed
intake is reduced, the added fat permits the hen to maintain egg formation while minimizing heat generated (Valencia et al.,
1980).
Fatty Acid Composition
Directly employing dietary fat in the assembly of either body or egg lipids results in a fatty acid composition similar to
that of the diet. Fat absorbed from the fowl's intestine is transported to the liver, where some modifications may occur. For
the most part, the unsaturated fatty acids are unchanged, but the saturated ones may undergo desaturation, especially stearic
acid which can be converted to oleic acid. Also, elongation and further desaturation of 18:2(n-6) and 18:3(n-3) may occur in
the liver.
Depot fat is the tissue most affected by the source of dietary fat. Depot fat of both broiler chickens (Schuler and Essary,
1971; Edwards et al., 1973) and turkeys (Moran et al., 1973; Salmon and O'Neil, 1973) are more influenced by the vegetable
oils having high proportions of polyunsaturated fatty acids than by more saturated animal fats.
Fatty acid composition in depots can be altered by changing from one dietary fat to another (Watkins, 1988). The extent
of influence that each fat has on body composition increases with the level of intake, duration of feeding, and stage of
maturity (Bartov et al., 1974; Salmon, 1976). The hen's adipose depots respond to dietary fat in the same way as do those of
growing birds, and the yolk lipid exhibits a fatty acid pattern resembling that of the dietary fat (Guenter et al., 1971; Sim et
al., 1973).
Essential Fatty Acids
Linoleic acid (18:2, n-6) and a-linolenic acid (18:3, n-3) are recognized as metabolically essential fatty acids. The
position of the double bonds in these n-6 and n-3 polyunsaturated fatty acids (PUFA) is unique because they are not formed
in the fowl. The essential fatty acids are converted to long-chain PUFA in poultry through a series of desaturation (addition of
a double bond) and elongation steps (chain-lengthening with 2 carbons) to form 20 and 22 carbon PUFA (Watkins, 1991).
Membrane phospholipids contain a greater proportion of PUFA than do triacyglyerols although depot fat can contain a
reserve of linoleic acid for the fowl. In poultry, specific PUFA are biosynthesized into compounds called eicosanoids which
act as potent biological regulators.
Linoleic acid is the only essential fatty acid for which a dietary requirement has been demonstrated. Inadequacies of
linoleic acid are not readily encountered, but symptoms that result are due to a loss of membrane integrity. An increased need
for water and decreased resistance to disease are characteristic deficiency symptoms observed in poultry (Balnave, 1970). A
deficiency of linoleic acid in the male can impair spermatogenesis and affect fertility. Insufficient deposition of linoleic acid
in the egg will adversely affect embryonic development. The essential fatty acid requirements of growing and adult birds can
usually be satisfied by feeding a diet with 1 percent of linoleic acid. Higher levels of linoleic acid may be needed by the
laying hen to achieve and maintain satisfactory egg weight.
A dietary need for α-linolenic acid (18:3, n-3) has yet to be demonstrated for the fowl. α-Linolenic acid appears to be
important, however, in the development of specialized membranes found in the retina and nervous system. These membranes
contain relatively high concentrations of n-3 PUFA that can originate from 18:3(n-3) (Neuringer and Connor, 1986).
Certain PUFA derived from linolenic and α-linolenic acids are biosynthesized into a multitude of eicosanoids. The
primary substrates for eicosanoid production are 20:4(n-6), 20:3(n-6) which are formed from linoleic acid, and 20:5(n-3) a
product of α-linolenic acid. Preceding eicosanoid biosynthesis in poultry, the PUFA is released from membrane
phospholipids by action of phospholipases. Liberation of PUFA is induced by a number of stimuli. Following a series of
different enzymatic steps, several eicosanoids can be formed depending on the tissue and cell type (Watkins, 1991). The
eicosanoids are categorized into prostaglandins, prostacyclins, thromboxanes, and leukotrienes. Formation of eicosanoids is
widespread in the body and nearly every physiological system is affected by these hormone-like compounds. The eicosanoids
are important in embryonic development, reproduction, immunological responses, and bone development in poultry
(Watkins, 1991).
Eicosanoid production can be modulated depending upon the concentration of substrate PUFA found in tissues.
Changing the dietary concentrations of n-3 and n-6 PUFA found in tissues will influence the types and amounts of
eicosanoids formed (Watkins, 1991). Elevating the n-3 PUFA content of the diet relative to that for n-6 PUFA alters
eicosanoid production in immunocompetent cells (Kinsella et al., 1990). These types of responses also seem to affect
inflammatory reactions and blood clotting in animals and humans. To maintain the full spectrum of eicosanoid effects in the
body a balanced intake of n-3 and n-6 PUFA is recommended.
MINERAL
Minerals are the inorganic part of feeds or tissues. They are often divided into two categories, based on the
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amount that is required in the diet. Requirements for major, or macro, minerals usually are stated as a percentage of the diet,
whereas requirements for minor, or trace, minerals are stated as milligrams per kilogram of diet or as parts per million.
Minerals are required for the formation of the skeleton, as components of various compounds with particular functions
within the body, as cofactors of enzymes, and for the maintenance of osmotic balance within the body of the bird. Calcium
and phosphorus are essential for the formation and maintenance of the skeleton. Sodium, potassium, magnesium, and chloride
function with phosphates and bicarbonate to maintain homeostasis of osmotic relationships and pH throughout the body.
Most of the calcium in the diet of the growing bird is used for bone formation, whereas in the mature laying fowl most dietary
calcium is used for eggshell formation. Other functions of calcium include roles in blood clotting and as a second messenger
in intracellular communications.
An excess of dietary calcium interferes with the availability of other minerals, such as phosphorus, magnesium,
manganese, and zinc. A ratio of approximately 2 calcium to 1 nonphytate phosphorus (weight/weight) is appropriate for most
poultry diets, with the exception of diets for birds that are laying eggs. When poultry are laying eggs, a much higher level of
calcium is needed for eggshell formation, and a ratio as high as 12 calcium to 1 nonphytate phosphorus (weight/weight) may
be correct. But high levels of calcium carbonate (limestone) and calcium phosphates may tend to make the diet unpalatable
and dilute the other dietary components. If a calcium source contains a high level of magnesium (as does dolomitic
limestone), it probably should not be used in poultry diets (Stillmak and Sunde, 1971).
Phosphorus, in addition to its function in bone formation, is also required in the utilization of energy and in structured
components of cells. Examples of phosphorus-containing compounds are adenosine 5'-triphosphate (ATP) and phospholipids.
These forms of phosphorus, if present in plants, can be digested by poultry; however, such digestible forms usually account
for only 30 to 40 percent of the total phosphorus. The remaining phosphorus is present as phytate phosphorus and is poorly
digested. Only about 10 percent of the phytate phosphorus in corn and wheat is digested by poultry (Nelson, 1976). The
phosphorus from animal products and phosphorus supplements is generally considered to be well utilized. Phosphorus
supplements for poultry diets are listed in Table 9-10.
Sodium and chloride are essential for all animals. Dietary concentrations of salt generally used are those that will just
support maximum growth rate or egg production. Higher concentrations lead to excessive consumption of water and
attendant problems with ventilation control and wet droppings.
Dietary proportions of sodium, potassium, and chloride are important determinants of acid-base balance (Mongin, 1968;
Hurwitz et al., 1973; Cohen and Hurwitz, 1974; Sauveur and Mongin, 1978). Other cations and anions such as calcium,
sulfate, and phosphate also may be involved. The appropriate dietary balance of these electrolytes is often assessed by the
levels of sodium and potassium versus chloride, where each element is expressed in milliequivalents per kilogram of diet.
Experiments show that sodium and potassium are alkalogenic (have an alkaline-producing effect), whereas chloride is
acidogenic (has an acid-producing effect). Chloride tends to decrease blood pH and bicarbonate concentration, whereas
sodium and potassium tend to increase blood pH and bicarbonate concentration. The proper dietary balance of sodium,
potassium, and chloride is necessary for growth, bone development, eggshell quality, and amino acid utilization (Mongin,
1981). However, an ideal balance among these electrolytes appropriate for a wide range of environmental situations has not
been defined.
Trace elements, including copper, iodine, iron, manganese, selenium, and zinc are required in small amounts in the diet.
Cobalt is also required, but it does not need to be supplied as a trace mineral because it is a part of vitamin B12. In practical
diets, copper and iron are often present at sufficient levels without supplementation.
Trace elements function as part of larger organic molecules. Iron is a part of hemoglobin and cytochromes, and iodine is
a part of thyroxine. Copper, manganese, selenium, and zinc function as essential accessory factors to enzymes and, in the case
of zinc, DNA structural motifs (zinc fingers). If one of these minerals is deficient, the functional activity of the organic
moiety requiring the presence of the mineral will be decreased, as has been described in detail for each mineral by Mertz
(1986).
The requirements for trace minerals are often fulfilled by concentrations present in conventional feed ingredients. Soils
vary, however, in their content of trace minerals, and plants vary in their uptake of minerals. Consequently, feedstuffs grown
in certain geographic areas may be marginal or deficient in specific elements. Thus, poultry diets may require
supplementation to ensure adequate intake of trace minerals. Because of the interactions that occur between various minerals
such as copper and molybdenum, selenium and mercury, calcium and zinc, calcium and manganese (Mertz, 1986), excessive
concentrations of one element may result in a deficiency in the amount available to the bird of some other element.
Formulators of poultry diets should be aware of these possible mineral interactions and of the
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potential effects that the chemical form (cation-anion combination) of mineral sources may have on their utilization by
poultry (Allaway, 1986). Mineral salts used as feed supplements are not usually pure compounds but contain variable
amounts of other minerals. The concentrations of minerals that may be present in feed-grade mineral supplements are shown
in Table 9-10.
Experimental diets may sometimes be formulated from purified or chemically defined ingredients. Under these
conditions, silicon and boron may be inadequate and biological responses may occur with the addition of these elements to
the diet (Carlisle, 1970, 1980; Nielsen, 1986).
VITAMINS
Vitamins are generally classified under two headings: fat soluble vitamins, A, D, E, and K, and water-soluble vitamins,
that include the so-called B-complex and vitamin C (ascorbic acid). Vitamin C is synthesized by poultry and is, accordingly,
not considered a required dietary nutrient. There is some evidence, nevertheless, of a favorable response to vitamin C by birds
under stress (Pardue et al., 1985).
The requirements for most vitamins are given in terms of milligrams per kilogram of diet. Exceptions are vitamins A, D,
and E, for which requirements are commonly stated in units. Units are used to express the requirements for these vitamins
because different forms of the vitamins have different biological activities (Anonymous, 1990).
Requirements for vitamin A are expressed in either International Units (IU) or U.S. Pharmacopeia units (USP) per
kilogram of diet. The international standards for vitamin A activity are as follows: 1 IU of vitamin A = 1 USP unit = vitamin
A activity of 0.3 µg crystalline vitamin A alcohol (retinol), 0.344 µg vitamin A acetate, or 0.55 µg vitamin A palmitate. One
IU of vitamin A activity is equivalent to the activity of 0.6 µg of β-carotene; alternatively, 1 mg β-carotene = 1,667 IU
vitamin A (for poultry).
Vitamin D for poultry must be in the form of vitamin D3, which is found naturally in fish liver oil or may be synthesized
by the irradiation of animal sterol. Vitamin D2, which is from plant sources, is active for rats and most mammals but has very
low activity for poultry. One unit of vitamin D3 (USP or IU) is defined as the activity of 0.025 µg of vitamin D3
(cholecalciferol). The requirements listed herein for vitamin D are based on diets containing the stated requirements for
calcium and available phosphorus.
One IU of vitamin E is the activity of 1 mg of synthetic DL-α-tocopheryl acetate, 0.735 mg D-α-tocopheryl acetate, 0.671
mg D-α-tocopherol, or 0.909 mg DL-α-tocopherol. The dietary requirement for vitamin E is highly variable and depends on the
concentration and type of fat in the diet, the concentration of selenium, and the presence of prooxidants and antioxidants.
Vitamin K activity is exhibited by a number of naturally occurring and synthetic compounds with varying solubilities in
fat and water. Menadione (2-methyl-1,4-naphthoquinone) is a fat soluble synthetic compound that can be considered the
reference standard for vitamin K activity. Two naturally occurring forms are K1 or phylloquinone (2-methyl-3-phytyl-1,4naphthoquinone) and K2 or menaquinone (K1 substituted with 2 to 7 isoprene units). Water-soluble forms include menadione
sodium bisulfite (MSB), menadione sodium bisulfite complex (MSBC), and menadione dimethylpyrimidol (MPB). The
theoretical activity of these compounds is 33, 50, and 45 percent, respectively, as calculated on the basis of the proportion of
menadione present in the molecule.
Dietary supplements frequently contain, as a factor of safety, levels of vitamins in considerable excess of the minimum
requirements. Vitamin tolerances have been reviewed by the National Research Council (1987b). Maximum tolerances for
vitamins are of the order of 10 to 30 times the minimum requirement for vitamin A, 4 to 10 times for vitamin D3, and 2 to 4
times for choline chloride (possibly because of the chloride). Niacin, riboflavin, and pantothenic acid are generally tolerated
at levels as great as 10- to 20-fold their nutritional requirement. Vitamin E is generally tolerated at intakes as great as 100fold the required level. Vitamins K and C, thiamin, and folic acid are generally tolerated at oral intake levels of at least 1,000fold the requirement. Pyridoxine may be tolerated at 50 times or more of the requirement (Aboaysha and Kratzer, 1979).
High levels of biotin and vitamin B12 have not been tested.
WATER
Water must be regarded as an essential nutrient, although it is not possible to state precise requirements. The amount
needed depends on environmental temperature and relative humidity, the composition of the diet, rate of growth or egg
production, and efficiency of kidney resorption of water in individual birds (Medway and Kare, 1959). It has been generally
assumed that birds drink approximately twice as much water as the amount of feed consumed on a weight basis, but water
intake actually varies greatly.
Several dietary factors influence water intake and water:feed ratios. Increasing crude protein increases water intake and
water:feed ratios (Marks and Pesti, 1984). Crumbling or pelleting of diets increases both water and
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16
feed intake relative to mash diets, but water:feed ratios stay relatively the same (Marks and Pesti, 1984). Increasing dietary
salt increases the water intake (Marks, 1987).
The data given for water consumption in Table 1-1 are for environmental temperatures of about 21°C except for
brooding chicks and poults. With broilers, water consumption increases about 7 percent for each 1°C above 21° C. Laying
hens may consume from 150 to 300 liters (40 to 80 gal) per 1,000 birds daily, depending on temperature and other factors.
Survival under extremely hot conditions is influenced by the ability to consume large quantities of water or, more precisely,
the ability to use water to remove heat from the respiratory surfaces of the body. This ability varies from strain to strain.
Water intake data for broilers listed herein are based on studies using modern commercial broilers (Marks, 1981; Ross
and Hurnik, 1983; Gardiner and Hunt, 1984; Pesti et al., 1985; Miller et al., 1988). Most of the studies were carried out under
moderate temperature conditions, with corrections for evaporative losses. In most of the studies, data also were collected on
feed intake, allowing for calculation of water:feed ratios.
Documented water intake data for laying hens are limited, especially data related to cage systems. Dun and Emmans
(1971) compared the water consumption of caged hens on trough and nipple watering systems in a 3-year study. Feed and
water consumption were 126 g and 254 ml with the trough system and 124.9 g and 166 ml with the nipple system (four hens
per nipple). Hearn and Hill (1978) compared feed and water consumption of hens on trough and nipple watering systems,
with varying numbers of birds per nipple. During the study, that was conducted from 20 to 72 weeks of age, hens on trough
waterers consumed an average of 115 g of feed and 213 ml of water. Hens with 2.5, 5, and 10 birds per nipple consumed 109,
109, and 108 g of feed and 182, 169, and 165 ml of water, respectively. Gardiner (1982) examined the water intake of
individually caged hens for a 336-day period beginning when they were 32 weeks of age. Over this period of time, mean feed
consumption of laying hens was 109 g and daily water intake was 183 ml, for a feed:water ratio of 1.68. There was no
indication of type of drinker used. It is evident that the type of watering system used will influence water consumption (or,
more correctly, water disappearance) of laying hens. Although many tables of estimated water consumption can be found in
the literature, the sources of the data used to compile these tables cannot be documented.
Water consumption data for turkeys obtained from experimental studies are meager (Enos et al., 1967). Thus, the data on
water consumption of turkeys shown in Table 1-1 are based mainly on information obtained recently from commercial turkey
production companies.
TABLE 1-1 Water Consumption by Chickens and Turkeys of Different Ages
Age (weeks)
Broiler Chickens (ml
White Leghorn
Brown-Egg-Laying
per bird per week)a
Hens (ml per bird
Hens (ml per bird
per week)a
per week)a
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
225
480
725
1,000
1,250
1,500
1,750
2,000
–
–
–
–
–
–
–
–
–
–
–
200
300
–
500
–
700
–
800
–
900
–
1,000
–
1,100
–
1,200
–
1,300
–
1,600
200
400
–
700
–
800
–
900
–
1,000
–
1,100
–
1,100
–
1,200
–
1,300
–
1,500
Large White Turkeys (ml per bird per week)
a,b
Males
385
750
1,135
1,650
2,240
2,870
3,460
4,020
4,670
5,345
5,850
6,220
6,480
6,680
6,800
6,920
6,960
7,000
7,020
7,040
Females
385
690
930
1,274
1,750
2,150
2,640
3,180
3,900
4,400
4,620
4,660
4,680
4,700
4,720
4,740
4,760
–
–
–
NOTE: Dash indicates that information is not available.
a Varies considerably depending on ambient temperature, diet composition, rates of growth or egg production, and type of equipment
used. The data presented apply under moderate (20° to 25°C) ambient temperatures.
b Based on data obtained from commercial turkey production units.
Water deprivation for 12 hours or more has adverse effects on the growth of young poultry and egg production of laying
hens, and water deprivation of 36 hours or more results in a marked increase in mortality of young and old poultry (Bierer et
al., 1965a,b; Haller and Sunde, 1966; Adams, 1973). Water restoration, after extended periods of water deprivation (36 to 40
hours), may cause a ''drunken syndrome" or "water intoxication," leading to death (Marsden et al., 1965). Young turkeys are
especially susceptible to this condition.
The salt content and pH of water may influence the use of the drinking water to administer vitamins and drugs. Turkeys
are known to detect minor differences in the flavor of medicated water and may accept drugs in one water supply but not in
another. Intermittent provision of water is sometimes used to reduce the water content of the droppings and to control feed
intake in laying hens without reducing egg production (Maxwell and Lyle, 1957). Because birds differ in their ability to
conserve body water by increasing kidney resorption, there is a danger of causing dehydration of some birds by practicing
water restriction of a flock.
Some water supplies contain considerable concentrations of sulfur or sulfates, nitrates, and various trace minerals. These
are usually readily absorbed from the intestine and may be either useful or harmful to the bird, depending
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on concentration. Table 1-2 gives the guidelines suggested by the National Research Council (1974) for the suitability for
poultry of water with different concentrations of total dissolved solids (TDS); that is, the total concentration of all dissolved
elements in water.
TABLE 1-2 Guidelines for Poultry for the Suitability of Water with Different Concentrations of Total Dissolved Solids (TDS)
Comments
TDS (ppm)
Less than 1,000
These waters should present no serious burden to any class of poultry.
1,000–2,999
These waters should be satisfactory for all classes of poultry. They may cause watery droppings (especially at the
higher levels) but should not affect health or performance.
3,000–4,999
These are poor waters for poultry, often causing watery droppings, increased mortality, and decreased growth
(especially in turkeys).
5,000–6,999
These are not acceptable waters for poultry and almost always cause some type of problem, especially at the upper
limits, where decreased growth and production or increased mortality probably will occur.
7,000–10,000
These waters are unfit for poultry but may be suitable for other livestock.
These waters should not be used for any livestock or poultry.
More than 10,000
SOURCE: National Research Council. 1974. Nutrients and Toxic Substances in Water for Livestock and Poultry. Washington, D.C.:
National Academy of Sciences.
XANTHOPHYLLS
A number of carotenoid pigments are responsible for the yellow-orange coloration of egg yolks and poultry fat and also
may contribute to coloration of the skin, shanks, feet, and beak. The xanthophylls, which are characterized by the presence of
hydroxyl groups, are the carotenoids of most interest in poultry nutrition. The most commonly considered xanthophylls are
lutein in forages such as alfalfa and zeaxanthin in corn. Relative xanthophyll contribution by various xanthophyll-rich
ingredients is shown in Table 1-3.
Individual xanthophylls differ in their ability to impart color. Although β-carotene has little pigmenting value, other
xanthophylls and synthetic products are effective in influencing yolk and skin color. Less than 1 percent of dietary β-carotene
is deposited in the yolk, but for zeaxanthin, as found in corn, the value is closer to 7 percent, and for some synthetic products,
such as β-apo-8-carotenoic acid ethyl ester, the incorporation rate may be as high as 34 percent (Roche Vitamins and Fine
Chemicals, 1988). Fletcher et al. (1985) and Saylor (1986) reported that natural sources of xanthophyll differed in their
ability to pigment egg yolk and the skin of broilers. Alfalfa meal contains several types of xanthophylls, but the one of
greatest abundance and importance is lutein, which tends to impart a yellow color, whereas corn and corn gluten meal contain
primarily zeaxanthin, which tends to impart an orange-red color.
TABLE 1-3 Xanthophyll and Lutein Content of Selected Ingredients
Ingredient
Alfalfa meal, 17% crude protein
Alfalfa meal, 22% crude protein
Alfalfa protein concentrate, 40% crude protein
Algae meal
Corn
Corn gluten meal, 60% crude protein
Marigold petal meal
Xanthophyll (mg/kg)
220
330
800
2,000
17
290
7,000
Lutein (mg/kg)
143
—
—
—
0.12
120
—
NOTE: Dash indicates that information is not available.
Avian tissue normally accumulates xanthophylls, although the retina may accumulate other carotenoids (Goodwin,
1986). In the laying hen, 50 percent of total body zeaxanthin (as derived from corn) is found in the ovary (Scheidt et al.,
1985). Goodwin (1986) indicated that body stores of xanthophylls in the muscle and skin are transferred to the ovary at onset
of sexual maturity. Presumably, this transfer occurs throughout the egg production cycle and contributes to the gradual loss of
pigment from the shank and beak as egg production continues.
Synthetic carotenoids that have been approved for use by regulatory agencies are used in poultry diets, because levels of
desired pigments in natural feedstuffs are not always constant and many of the carotenoid-containing natural feedstuffs are
relatively low in energy content. Approval of use of these synthetics varies among countries. Synthetic pigments, such as
canthaxanthin and β-apo-8-carotenoic acid (usually as an ethyl ester), can be used to control pigmentation more precisely to
yield varying degrees of yellow-orange-red coloration. In natural products, xanthophylls are unstable, and effective levels
may decline as a result of oxidation during prolonged storage. This decline can be reduced by the inclusion of antioxidants in
the feed.
A number of factors can adversely affect absorption of xanthophylls and thus lead to reduced pigmentation. Broilers
infected with Eimeria sp. exhibit reduced pigmentation and blood xanthophylls (Bletner et al., 1966), and the viral infection
that may be responsible for malabsorption syndrome also results in altered xanthophyll status of the bird (Winstead et al.,
1985). Exposing feed to light may have variable effects on subsequent pigmentation (Fletcher, 1981). The presence of certain
mycotoxins in feeds seems to be detrimental to pigmentation (Tyczkowski and Hamilton, 1987).
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COMPONENTS OF POULTRY DIETS
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UNIDENTIFIED GROWTH FACTORS
So-called unidentified growth factors have been reported throughout the history of poultry nutrition studies. Natural
ingredients claimed to contain such factors are most often animal proteins or fermentation by-products (Summers et al., 1959;
Al-Ubaidi and Bird, 1964; Dixon and Couch, 1970; Waldroup et al., 1970). Ingredients containing unidentified growth
factors are claimed to improve chick growth and reproductive performance (Morrison et al., 1956; Touchburn et al., 1972).
Bhargava and Sunde (1969) described a chick assay for quantitation of such unidentified factors.
The mode of action of these unidentified factors is far from clear, however. With the identification of vitamins and
consideration of the significance of trace minerals, many nutritionists now disregard the importance of growth factors. That
responses may still occur could relate to truly unidentified nutrients or, more likely, to changes in feed palatability and/or
quality (Alenier and Combs, 1981; Cantor and Johnson, 1983), mineral chelation, or simple improvement in the balance of
available nutrients.
ANTIMICROBIALS
Antimicrobial feed additives, although not nutrients in the sense that they are required by poultry, are included in diets to
improve growth, efficiency of feed utilization and livability (Stokstad et al., 1949; Coates et al., 1951; Libby and Schaible,
1955; Milligan et al., 1955; Bird, 1968; Begin, 1971; Morrison et al., 1974). Antimicrobial agents are included in diets at
relatively low concentrations (1 to 50 mg/kg), depending on the agent and stage of development of poultry. They are,
accordingly, classified as additives and as growth promoters. Egg production is also frequently improved by dietary
supplementation with antimicrobial agents (Carlson et al., 1953; Balloun, 1954; Andrews et al., 1966). The mechanisms by
which antimicrobials improve performance are not clearly understood. Because antimicrobials do not stimulate growth of
chicks kept in a germfree environment (Coates and Harrison, 1969), it is likely that stimulation of growth results from either
suppression of microorganisms that may cause adverse effects or encouragement of other microorganisms that may have
favorable effects on poultry performance.
There is some concern that feeding of low concentrations of antibiotics may favor the proliferation of antibiotic-resistant
microorganisms, which could have serious consequences for disease control in humans or domestic animals. A study by the
National Research Council (1980a) examined this concern and concluded that "the postulations concerning the hazards to
human health that might result from the addition of subtherapeutic antimicrobials to feeds have been neither proven nor
disproven." Continued monitoring of bacterial resistance in humans and animals has not provided clear-cut answers to this
concern.
Constraints and regulations on use of particular antimicrobials in poultry feeds vary among countries and are subject to
change. Detailed information on specific antimicrobial agents, levels of usage, and legal requirements for use in the United
States and Canada may be found in the Feed Additive Compendium (published each year by the Miller Publishing Company,
2501 Wayzata Boulevard, Minneapolis, MN 55440) and in the compendium of "Medicating Ingredient Brochures" (Plant
Products Division, Canada Department of Agriculture, Ottawa, Ontario, Canada).
For official information concerning Food and Drug Administration approval of antibiotics and other animal drugs, the
Code of Federal Regulations (CFR), Title 21, should be consulted. Title 21 is revised at least once each year as of April 1.
The CFR is kept up to date by the individual issues of the Federal Register. These two publications must be used together to
determine the latest version of any given rule. Title 21 is published in six parts: Part 500-599 covers animal drugs, feeds, and
related products and is available from the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C.
20402. The Federal Register is available from the Superintendent of Documents and includes monthly issues of the "List of
CFR Sections Affected" and "The Federal Register Index."
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NUTRIENT REQUIREMENTS OF CHICKENS
19
2
Nutrient Requirements of Chickens
Chickens vary greatly according to the purpose for which they have been developed. Those intended for the production
of eggs for human consumption (Leghorn-type) have a small body size and are prolific layers, whereas those used as broilers
or broiler breeders (meat-type) have rapid growth rates and a large body size. They are less efficient egg layers. Methods of
feeding differ for these two kinds of chickens.
LEGHORN-TYPE CHICKENS
Methods of feeding Leghorn-type chickens depend on the age and activity (laying or breeding) of the bird. Feed
requirements change as birds pass through the starting and growing, pre-egg-laying, egg production, and molt phases.
Starting and Growing Pullets
Relatively little research has been conducted in the last 10 years to obtain definitive nutrient requirements for immature
Leghorn-type birds. In large part, this situation is due to the use of meat-strain birds in requirement studies involving avian
species. Thus, although growth and maturity characteristics of egg-strain pullets have changed considerably over the last 10
years, particularly for brown-egg-laying birds, the only data available on requirements for many nutrients are dated. Most
current research activity deals with nutrients of major economic significance. The available information is reviewed in
Appendix Table A-1.
Nutrient requirements of immature Leghorn-type chickens (pullets) are listed in Table 2-1. Although requirements are
assessed ultimately in terms of subsequent reproductive performance, the criteria used by the committee were adequate
growth rate (in terms of final body weight at different ages) and normal metabolism. It is well documented that mature body
weight can greatly influence the subsequent reproductive performance (Leeson and Summers, 1987a), and, as such, this
criterion becomes critical in the assessment of nutritional status.
The dearth of research information for immature pullets is even more acute for brown-egg-laying strains. Because
brown-egg-laying birds predominate in many parts of the world, the committee has attempted to define their nutrient
requirements as well. In large part, however, these requirement values have been extrapolated from studies conducted with
Leghorns with consideration for the larger body weight and/or appetite and increased maintenance requirement of brown-egg
layers.
The nutrient requirement values shown in Table 2-1 and the performance characteristics shown in Table 2-2 are based
on the assumption that the birds will be allowed to consume feed in an ad libitum manner. Ad libitum feed consumption is
important for Leghorn birds, especially when reared in hot climates, because of their inherently low appetites. Managers
should routinely consider restricted feeding only for brown-egg-laying strains, and even then only in temperate climates and
with high-energy diets.
Protein And Energy
In discussing the protein needs of growing pullets, it is assumed that the amino acid profile is balanced according to the
requirement values shown in Table 2-1. Pullets allowed to self-select diets based on protein or energy content seem to
voluntarily consume much less protein in early life and more protein as they approach maturity (Summers and Leeson, 1978)
than do pullets on more conventional programs. However, low-protein or low-lysine starter diets invariably depress the growth
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NUTRIENT REQUIREMENTS OF CHICKENS
TABLE 2-1 Nutrient Requirements of Immature Leghorn-Type Chickens as Percentages or Units per Kilogram of Diet
White-Egg-Laying Strains
Brown-Egg-Laying Strains
Nutrient
Unit
0 to 6
6 to 12
12 to 18
18
0 to 6
6 to 12
12 to 18
Weeks;
Weeks;
Weeks;
Weeks;
Weeks
Weeks;
Weeks;
a
a
a
a
a
500 g
450 g
980 g
1,375 g
to First
1,100 g
1,500 ga
Egg;
2,850b
2,850b
2,900b
2,800b
2,800b
2,850b
1,475 ga
2,900b
Protein and
amino acids
Crude proteinc
%
18.00
16.00
15.00
17.00
17.00
15.00
14.00
Arginine
%
1.00
0.83
0.67
0.75
0.94
0.78
0.62
Glycine + serine
%
0.70
0.58
0.47
0.53
0.66
0.54
0.44
Histidine
%
0.26
0.22
0.17
0.20
0.25
0.21
0.16
Isoleucine
%
0.60
0.50
0.40
0.45
0.57
0.47
0.37
Leucine
%
1.10
0.85
0.70
0.80
1.00
0.80
0.65
Lysine
%
0.85
0.60
0.45
0.52
0.80
0.56
0.42
Methionine
%
0.30
0.25
0.20
0.22
0.28
0.23
0.19
%
0.62
0.52
0.42
0.47
0.59
0.49
0.39
Methionine +
cystine
Phenylalanine
%
0.54
0.45
0.36
0.40
0.51
0.42
0.34
Phenylalanine +
%
1.00
0.83
0.67
0.75
0.94
0.78
0.63
tyrosine
Threonine
%
0.68
0.57
0.37
0.47
0.64
0.53
0.35
Tryptophan
%
0.17
0.14
0.11
0.12
0.16
0.13
0.10
Valine
%
0.62
0.52
0.41
0.46
0.59
0.49
0.38
Fat
Linoleic acid
%
1.00
1.00
1.00
1.00
1.00
1.00
1.00
Macrominerals
%
0.90
0.80
0.80
2.00
0.90
0.80
0.80
Calciumd
Nonphytate
%
0.40
0.35
0.30
0.32
0.40
0.35
0.30
phosphorus
Potassium
%
0.25
0.25
0.25
0.25
0.25
0.25
0.25
Sodium
%
0.15
0.15
0.15
0.15
0.15
0.15
0.15
Chlorine
%
0.15
0.12
0.12
0.15
0.12
0.11
0.11
Magnesium
mg
600.0
500.0
400.0
400.0
570.0
470.0
370.0
Trace minerals
Manganese
mg
60.0
30.0
30.0
30.0
56.0
28.0
28.0
Zinc
mg
40.0
35.0
35.0
35.0
38.0
33.0
33.0
Iron
mg
80.0
60.0
60.0
60.0
75.0
56.0
56.0
Copper
mg
5.0
4.0
4.0
4.0
5.0
4.0
4.0
Iodine
mg
0.35
0.35
0.35
0.35
0.33
0.33
0.33
Selenium
mg
0.15
0.10
0.10
0.10
0.14
0.10
0.10
Fat soluble
vitamins
A
IU
1,500.0
1,500.0
1,500.0
1,500.0
1,420.0
1,420.0
1,420.0
ICU
200.0
200.0
200.0
300.0
190.0
190.0
190.0
D3
E
IU
10.0
5.0
5.0
5.0
9.5
4.7
4.7
K
mg
0.5
0.5
0.5
0.5
0.47
0.47
0.47
Water soluble
vitamins
Riboflavin
mg
3.6
1.8
1.8
2.2
3.4
1.7
1.7
Pantothenic acid
mg
10.0
10.0
10.0
10.0
9.4
9.4
9.4
Niacin
mg
27.0
11.0
11.0
11.0
26.0
10.3
10.3
mg
0.009
0.003
0.003
0.004
0.009
0.003
0.003
B12
Choline
mg
1,300.0
900.0
500.0
500.0
1,225.0
850.0
470.0
Biotin
mg
0.15
0.10
0.10
0.10
0.14
0.09
0.09
Folic acid
mg
0.55
0.25
0.25
0.25
0.52
0.23
0.23
Thiamin
mg
1.0
1.0
0.8
0.8
1.0
1.0
0.8
mg
3.0
3.0
3.0
3.0
2.8
2.8
2.8
Pyridoxine
20
18
Weeks
to First
Egg;
1,600 ga
2,850b
16.00
0.72
0.50
0.18
0.42
0.75
0.49
0.21
0.44
0.38
0.70
0.44
0.11
0.43
1.00
1.80
0.35
0.25
0.15
0.11
370.0
28.0
33.0
56.0
4.0
0.33
0.10
1,420.0
280.0
4.7
0.47
1.7
9.4
10.3
0.003
470.0
0.09
0.23
0.8
2.8
NOTE: Where experimental data are lacking, values typeset in bold italics represent an estimate based on values obtained for other ages
or related species.
a Final body weight.
b These are typical dietary energy concentrations for diets based mainly on corn and soybean meal, expressed in kcal ME /kg diet.
n
c Chickens do not have a requirement for crude protein per se. There, however, should be sufficient crude protein to ensure an adequate
nitrogen supply for synthesis of nonessential amino acids. Suggested requirements for crude protein are typical of those derived with
corn-soybean meal diets, and levels can be reduced somewhat when synthetic amino acids are used.
d The calcium requirement may be increased when diets contain high levels of phytate phosphorus (Nelson, 1984).
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NUTRIENT REQUIREMENTS OF CHICKENS
21
rate of both white-egg- (Douglas and Harms, 1982; Kwakkel et al., 1991) and brown-egg-laying pullets (Maurice et al.,
1982), and early growth depression often depresses mature body weight and thereby adversely affects adult performance
(Milby and Sherwood, 1953; Leeson and Summers, 1979, 1987a). Low-protein diets have a transitory effect on muscle fiber
size rather than any long-term effect on numbers of such fibers (Timson et al., 1983). Although low-protein diets seem to
adversely affect growth rate, there is little indication that excessively high levels of protein have any benefit on growth and
development. Data of Keshavarz (1984) and Leeson and Summers (1989) suggest that in Leghorn pullets reduction in growth
is often seen when total protein intake to 140 days of age is less than 1 kg. An intake of 1 kg of balanced protein during the
same period seems to result in maximum growth.
TABLE 2-2 Body Weight and Feed Consumption of Immature Leghorn-Type Chickens
White-Egg-Laying Strains
Brown-Egg-Laying Strains
Age (weeks)
Body Weighta (g)
Feed Consumption (g/week)
Body Weighta (g)
Feed Consumption (g/week)
0
35
50
37
70
2
100
140
120
160
4
260
260
325
280
6
450
340
500
350
8
660
360
750
380
10
750
380
900
400
12
980
400
1,100
420
14
1,100
420
1,240
450
16
1,220
430
1,380
470
18
1,375
450
1,500
500
1,475
500
1,600
550
20
a Average genetic potential when feed is consumed on an ad libitum basis.
Different commercial strains may show different growth rates and different final mature body weights.
Energy intake may be the limiting factor for growth of egg-strain birds reared under most environmental conditions.
Assuming no amino acid deficiency, and an intake of 1 kg of protein from 1 day to 20 weeks, growth and development seem
most responsive to energy intake (Leeson and Summers, 1989). A total intake of 21 Mcal ME to 20 weeks seems ideal for
white-egg-laying pullets. However, manipulation of energy intake is not always easy, since the pullet appears to have a fairly
precise innate ability to regulate its energy intake regardless of dietary energy level (Cunningham and Morrison, 1976;
McNaughton et al., 1977b; Doran et al., 1983). Manipulation of energy intake is, therefore, best considered in relation to
feeding management and, in particular, methods of stimulating feed intake. For example, feed intake may be increased
through use of pelleted feed, increased frequency of feeding, feeding at cooler times of the day, and, where possible, use of
longer periods of light. Leeson and Summers (1989) concluded that pullet growth is initially most sensitive to dietary protein
and amino acids, whereas energy intake becomes more critical as the bird approaches maturity.
Skeletal size has also been considered as a criterion for assessment of pullet development. Lerner (1946) suggested that
skeletal size is a limiting factor for growth, and Jaap (1938) indicated that shank length can be used as a reliable estimate of
skeletal size per se. Skeletal development is related to adequate supplies of calcium, phosphorus, and vitamin D3, although
deficiencies of most nutrients can adversely affect normal vascularization of cartilage at the growth plate, a prerequisite to
normal calcification (Leeson and Summers, 1988). Skeletal growth is intimately associated with general growth and
development, and it is difficult to influence either independently. Leeson and Summers (1984) indicated that increased
skeletal size of pullets in response to dietary protein was associated with reduced ash content of bones.
Minerals And Vitamins
As indicated above, little work has been done recently to evaluate the mineral and vitamin requirements of young eggstrain birds. There has been some interest in reevaluating nonphytate phosphorus needs, although, in general, the new data
indicate no major change in previously reported requirement values. Both the young white-egg- (Douglas and Harms, 1986)
and the young brown-egg-laying pullets (Carew and Foss, 1980) exhibit an inferior growth rate when fed starter diets
containing less than 0.4 percent nonphytate phosphorus. The sodium requirement of the Leghorn pullet is approximately 0.15
percent of the diet regardless of age, although somewhat lower levels can be used after 10 weeks of age if excessive water
intake is problematic (Manning and McGinnis, 1980).
Prelay Period
Daily nutrient requirements of pullets 10 to 17 days before first egg are generally considered to be greater than during
the preceding 4 to 6 week period, although there is little evidence to show that pullets cannot meet these requirements
through increased voluntary feed intake.
Hoyle and Garlich (1987) found no change in growth or development of Leghorn pullets in response to elevated levels
of dietary energy or protein. As suggested above, energy intake is probably the most critical component for this age of bird,
and energy intake can perhaps be manipulated best through stimulation of feed intake rather than by simply increasing the
energy level of the feed.
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NUTRIENT REQUIREMENTS OF CHICKENS
22
The committee's review of research on the changes in metabolism of medullary bone immediately prior to maturity has
led to reevaluation of the pullets' requirement for calcium at this time. Since modern egg-strain pullets exhibit a rapid increase
in egg production and prolonged first multiegg clutch, it is obvious that a change in the requirements related to calcification
must be accommodated before or at time of first egg. Keshavarz (1987) indicated that feeding a diet containing 3.5 percent
calcium from as early as 14 weeks of age had no adverse effect on skeletal integrity, apparent renal function, or subsequent
reproductive performance. Leeson et al. (1986, 1987a) also observed normal pullet development, skeletal integrity, and
kidney histology when immature 19-week-old pullets were fed diets containing 3.5 percent calcium. These same workers
indicated that calcium levels of 0.9 to 1.5 percent at this age were detrimental to early shell quality. In studies in which
pullets were allowed to self-select nutrients, Classen and Scott (1982) showed that the birds consumed calcium in relation to
needs for deposition of medullary bone and (or) onset of shell calcification.
There has been little research on the phosphorus and vitamin D3 requirements of the prelay pullet.
Hens in Egg Production
Progress continues in the quest to use less feed in producing eggs. Most of this progress has resulted from decreasing the
amount of feed that is required for body maintenance of laying hens.
Body Maintenance Needs
Management practices, as well as nutritional regimes, can affect the maintenance requirement. In warmer houses, layers
need less energy from their feed because they expend less energy in maintaining body temperature. Hens eat less feed with
increasing temperatures and decrease feed consumption drastically at temperatures above 30°C (Davis et al., 1973; National
Research Council, 1981c).
Genetic selection can also affect the amount of feed required for maintenance. With chickens bred for higher rates of egg
production, there is a decrease in the maintenance requirement relative to eggs produced. At a rate of 100 percent egg
production (that is, one egg per hen per day), maintenance requirements must be fulfilled for the 12 days needed to produce a
dozen eggs; at a rate of 75 percent egg production, 16 days of maintenance requirements must be met to obtain a dozen eggs.
Body size also affects maintenance requirements. A compilation of information from nonpasserine birds showed that
basal metabolism was equal to 78.3 kcal per day × (kg body weight)723 (Lasiewski and Dawson, 1967). Conditions for
collection of these data were that the birds were in a postabsorptive state, in a thermoneutral environment, and as nearly at
rest as possible. Maintenance requirement, or the energy needed to sustain normal body processes and activities other than
growth and egg production, is greater than that of basal metabolism. In the thermoneutral range of temperatures, maintenance
for hens is approximately 100 kcal per day per kg body weight (MacLeod and Jewitt, 1988; Pesti et al., 1990). Strains of hens
may differ in their maintenance needs because of metabolic or behavioral characteristics (Pesti et al., 1990).
Production Needs
Nutritional factors can affect the amount of feed required to produce eggs. For example, some research indicates that
hens are able to make a good adjustment of feed intake to provide nearly identical daily energy intakes with up to 6 percent
added dietary fat (Sell et al., 1987). But other research suggests that the hen is not very accurate in adjusting feed intake to
provide equal daily energy intake when offered a range of dietary energy conditions (Morris, 1968; Rising et al., 1989).
Regardless of the accuracy of energy adjustment, hens eat less of a high-energy, nutritionally balanced feed than of a lowenergy feed to produce a dozen eggs.
Now that eggs can be produced with less feed, nutritionists have been permitted, or sometimes forced, to formulate diets
differently than they did several years ago. Generally, it is assumed that a hen's daily requirements for nutrients, other than
energy, are not changed by the level of feed consumption. If this is correct, then the difference in composition between the
diet of a layer eating 80 g of feed per day and the diet of one eating 120 g of feed per day should be about 40 g of energysupplying ingredients. But differences in daily feed consumption can cause the need for dramatic differences in dietary
nutrient concentration, if diets are formulated to supply a specified amount of nutrient, other than energy, each day. Nutrient
requirements of egg-type laying hens (Table 2-3) are expressed in terms of dietary concentrations for three levels of daily
feed consumption. (The research reports on which the committee based its nutrient requirement decisions are listed in
Appendix Table A-2.) Just how different rates of feed consumption can influence the formulation of a diet can be seen by
using one nutrient—say, lysine, as an example. The lysine required each day by a white-egg-laying hen is 690 mg, or 0.69 g.
Thus the diet of a white-egg-laying layer eating 100 g of feed per day should have a lysine concentration of 0.69 percent.
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NUTRIENT REQUIREMENTS OF CHICKENS
TABLE 2-3 Nutrient Requirements of Leghorn-Type Laying Hens as Percentages or Units per Kilogram of Diet (90 percent dry matter)
Dietary
Amounts Required per Hen Daily (mg or IU)
Concentrations
White-Egg
White-Egg
Brown-Egg
Required by WhiteBreeders at
Layers at
Layers at 110
Egg Layers at
100 g of Feed
100 g of
g of Feed per
Different Feed Intakes
per Hen Dailyb
Feed per Hen
Hen Dailyc
Daily
Nutrient
Unit
80a,b
100a,b
120a,b
Protein and amino
acids
%
18.8
15.0
12.5
15,000
15,000
16,500
Crude proteind
%
0.88
0.70
0.58
700
700
770
Argininee
Histidine
%
0.21
0.17
0.14
170
170
190
Isoleucine
%
0.81
0.65
0.54
650
650
715
Leucine
%
1.03
0.82
0.68
820
820
900
Lysine
%
0.86
0.69
0.58
690
690
760
Methionine
%
0.38
0.30
0.25
300
300
330
Methionine + cystine
%
0.73
0.58
0.48
580
580
645
Phenylalanine
%
0.59
0.47
0.39
470
470
520
%
1.04
0.83
0.69
830
830
910
Phenylalanine +
tyrosine
Threonine
%
0.59
0.47
0.39
470
470
520
Tryptophan
%
0.20
0.16
0.13
160
160
175
Valine
%
0.88
0.70
0.58
700
700
770
Fat
Linoleic acid
%
1.25
1.0
0.83
1,000
1,000
1,100
Macrominerals
f
%
4.06
3.25
2.71
3,250
3,250
3,600
Calcium
Chloride
%
0.16
0.13
0.11
130
130
145
Magnesium
mg
625
500
420
50
50
55
Nonphytate
%
0.31
0.25
0.21
250
250
275
phosphorusg
Potassium
%
0.19
0.15
0.13
150
150
165
Sodium
%
0.19
0.15
0.13
150
150
165
Trace minerals
Copper
mg
?
?
?
?
?
?
Iodine
mg
0.044
0.035
0.029
0.010
0.004
0.004
Iron
mg
56
45
38
6.0
4.5
5.0
Manganese
mg
25
20
17
2.0
2.0
2.2
Selenium
mg
0.08
0.06
0.05
0.006
0.006
0.006
Zinc
mg
44
35
29
4.5
3.5
3.9
Fat soluble vitamins
A
IU
3,750
3,000
2,500
300
300
330
IU
375
300
250
30
30
33
D3
E
IU
6
5
4
1.0
0.5
0.55
K
mg
0.6
0.5
0.4
0.1
0.05
0.055
Water soluble
vitamins
mg
0.004
0.004
0.004
0.008
0.0004
0.0004
B12
Biotin
mg
0.13
0.10
0.08
0.01
0.01
0.011
Choline
mg
1,310
1,050
875
105
105
115
Folacin
mg
0.31
0.25
0.21
0.035
0.025
0.028
Niacin
mg
12.5
10.0
8.3
1.0
1.0
1.1
Pantothenic acid
mg
2.5
2.0
1.7
0.7
0.20
0.22
Pyridoxine
mg
3.1
2.5
2.1
0.45
0.25
0.28
Riboflavin
mg
3.1
2.5
2.1
0.36
0.25
0.28
mg
0.88
0.70
0.60
0.07
0.07
0.08
Thiamin
NOTE: Where experimental data are lacking, values typeset in bold italics represent an estimate based on values obtained for other ages
or related species.
a Grams feed intake per hen daily.
b Based on dietary ME concentrations of approximately 2,900 kcal/kg and an assumed rate of egg production of 90 percent (90 eggs per
n
100 hens daily).
c Italicized values are based on those from white-egg layers but were increased 10 percent because of larger body weight and possibly
more egg mass per day.
d Laying hens do not have a requirement for crude protein per se. However, there should be sufficient crude protein to ensure an adequate
supply of nonessential amino acids. Suggested requirements for crude protein are typical of those derived with corn-soybean meal diets,
and levels can be reduced somewhat when synthetic amino acids are used.
e Italicized amino acid values for white-egg-laying chickens were estimated by using Model B (Hurwitz and Bornstein, 1973), assuming
a body weight of 1,800 g and 47 g of egg mass per day.
f The requirement may be higher for maximum eggshell thickness.
g The requirement may be higher in very hot temperatures.
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23
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NUTRIENT REQUIREMENTS OF CHICKENS
24
Hens eating 80 g of feed per day need a dietary lysine concentration of 0.86 percent to obtain 0.69 g per day; hens eating
120 g per day need a dietary lysine concentration of only 0.58 percent lysine to provide 0.69 g per hen per day. The basic
concept is that high daily feed consumption permits low nutrient concentrations and low daily feed consumption demands
high nutrient concentrations.
Equations have been developed to predict the energy required by chickens during egg production (McDonald, 1978;
National Research Council, 1981c). These equations use the expected energy requirements of hens as related to body weight,
daily egg mass, change in body weight, and ambient temperature to predict a total daily energy requirement. The data of
Table 2-4 show the predicted daily energy requirements of hens as related to different body weights and rates of egg
production, assuming no change in body weight and an ambient temperature of 22°C. The energy requirements derived from
such calculations can be used to estimate daily feed intake by relating the hen's energy needs to the dietary energy
concentration. Diets for laying hens, however, can be most accurately formulated on the basis of feed intake data obtained
frequently (every 1 to 2 weeks) for individual flocks.
Most egg-type hens are given ad libitum access to feed; however, feeding programs may be modified after the maximum
rate of egg mass output has been attained (Cerniglia et al., 1984; Cunningham, 1984). Laying hens eat more feed than is
needed to support egg production. As a result, it may be more profitable to limit their feed intake. Doing so would also reduce
the likelihood of health problems that can also result when hens are overly fat. Data on feed consumption in individual flocks,
together with information on body weight, ambient temperature, and rate of egg production, may be used to determine the
degree of feed restriction deemed appropriate.
Phase Feeding
Nutrient requirements presented in Table 2-3 assume that the amount of nutrient needed each day remains the same
throughout a hen's time of production. Some feeding programs, however, are based on the assumption that the amount of
nutrient needed each day is different at different stages of the production cycle. These programs are called phase feeding.
In phase feeding for flocks of laying hens, Phase 1 is designated as the time from the onset of egg production until past
the time of the maximum egg mass output, usually at about 36 weeks of age, which is the time of maximum egg mass output.
Phase 2 is the period between 36 and approximately 52 weeks, a period of high but declining egg production and increasing
egg weight. Phase 3 is from about 52 weeks to the end of the production cycle, in some instances to 80 weeks. During Phase
3 the rate of egg production continues to decline while egg weight increases only slightly.
TABLE 2-4 Estimates of Metabolizable Energy Required per Hen per Day by Chickens in Relation to Body Weight and Egg Production
(kcal)
Rate of Egg Production (%)
Body
Weight (kg)
0
50
60
70
80
90
1.0
130
192
205
217
229
242
1.5
177
239
251
264
276
289
2.0
218
280
292
305
317
330
2.5
259
321
333
346
358
371
296
358
370
383
395
408
3.0
NOTE: A number of formulas have been suggested for prediction of the daily energy requirements of chickens. The formula used here
was derived from that in Effect of Environment on Nutrient Requirements of Domestic Animals (National Research Council, 1981c):
ME per hen daily = W0.75 (173 - 1.95T) + 5.5 δW + 2.07 EE
where W = body weight (kg), T = ambient temperature (°C), δW = change in body weight (g/day), and EE = daily egg mass (g).
Temperature of 22°C, egg weight of 60 g, and no change in body weight were used in calculations.
A phase feeding program adjusts daily nutrient intakes according to expected requirements for maintenance and egg
production. Generally, daily intakes of protein, amino acids, and phosphorus are reduced with each succeeding phase. Daily
calcium intake usually is increased with each phase. Thus the dietary concentrations of these nutrients are changed
accordingly.
The scientific validity of the phase feeding concept has not been established. Experimental results have failed to prove
that a hen requires more nutrient per day at one stage of production than at another stage (Latshaw, 1981; Ousterhout, 1981;
Sell et al., 1987). Relatively low levels of feed intake during early egg production, however, necessitate the use of high
nutrient concentrations in diets during this phase of production.
Egg Weight
Egg weight is correlated with body weight of laying hens (Jull, 1924). The relative egg weight during a laying cycle
parallels the relative body weight. Within a flock, heavier birds lay heavier eggs (Leeson and Summers, 1987a). A body
weight decline in summer may account for the production of smaller eggs during that season (Cunningham et al., 1960).
Nutritional means may be used to alter egg weight slightly. Early in the egg production cycle, the objective would be to
increase egg weight. In one study (Summers and Leeson, 1983), the weight of eggs from pullets was not affected by increases
in dietary levels of methionine, linoleic acid, or protein above the established requirement. Another study showed that
increasing the level of dietary linoleic acid from 0.6 percent to 4.3 percent increased by egg weight during the first 14 weeks
of production; however, average daily egg yield was not affected (March and MacMillan, 1990). In a different study, adding
3 or 6 percent fat to diets fed during early
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NUTRIENT REQUIREMENTS OF CHICKENS
25
egg production increased egg weight by increasing yolk weight whether the diets were isocaloric or nonisocaloric (Sell et al.,
1987).
When egg weight is increased by fat supplementation of diets, it is not known if the response is due to fat in general or is
a specific response to linoleic acid (Whitehead, 1981; Balnave, 1982; Scragg et al., 1987). Increasing the percentage of fat or
oil in isoenergetic diets caused hens to lay heavier eggs (Whitehead, 1981; Sell et al., 1987). Decreasing the dietary energy
level, as may occur when sorghum or barley is substituted for corn, may decrease egg weight (Coon et al., 1988). Diet costs
may increase when supplemental fats are used to obtain higher dietary fat and energy concentrations. Thus managers should
determine the economic effectiveness of increasing egg weight in this way.
Older laying hens produce a high proportion of extralarge eggs for which monetary returns often do not offset costs of
production. Thus, a goal of feed formulators may be to reduce the weight of eggs produced by older hens. Decreasing dietary
levels of the most limiting amino acid can affect egg weight (Morris and Gous, 1988). For example, weight of eggs produced
by hens more than 38 weeks of age was reduced by limiting methionine intake to 270 mg per hen daily, compared with
feeding 300 mg methionine per hen daily (Peterson et al., 1983). A review of 12 scientific papers indicated that as the most
limiting amino acid level decreased below the required level, egg weight and rate of egg production were proportionally
reduced. This reduction occurred until egg weight decreased to about 90 percent of maximum. Further decreases in the amino
acid level decreased only the rate of egg production. An exception to the general effects of amino acid adequacy and egg
weight occurs with tryptophan, whereby a deficiency of this amino acid failed to decrease egg weight (Jensen et al., 1990).
Minerals And Vitamins
Mineral requirements of egg-type chickens in production are similar to mineral requirements of other poultry, with the
exception of calcium. The onset of egg production creates a need for more calcium to make the eggshell.
A question arises about the best time to switch pullets from a low-calcium growing diet to a high-calcium laying diet.
Feeding a diet with 3.25 percent calcium starting at 50 days of age increased the incidence of urolithiasis in later life
(Wideman et al., 1985). Changing from a low- to a high-calcium diet at 14 weeks of age or later, however, caused no
detrimental effects on performance through 60 weeks (Keshavarz, 1987). Although high-calcium levels are detrimental when
fed early in a pullet's life, feeding high-calcium levels several weeks before the onset of egg production seems to do no harm.
The calcium requirement listed in Table 2-3 is similar to values listed in earlier editions. Definitive research is still
lacking regarding several questions, however. Tests that cover a whole production cycle and that provide increments of
calcium ranging from 3 to 4.5 g per hen daily would be helpful. Such tests would answer questions related to amounts of
calcium needed, especially for the maintenance of eggshell strength in older layers. Conditions under which larger-particlesize calcium sources consistently improve eggshell strength should also be identified.
Levels of nutrients other than calcium may also affect eggshell strength. A wide sodium-to-chloride ratio can increase
blood pH and bicarbonate concentrations (Cohen et al., 1972). These increases may be the mechanism by which eggshell
strength is improved at thermoneutral zone temperatures with some diets when sodium chloride is replaced by sodium
bicarbonate in the water (Frank and Burger, 1965) or feed (Miles and Harms, 1982; Makled and Charles, 1987).
Phosphorus levels may also affect eggshell strength. Excess dietary phosphorus may decrease eggshell strength (Arscott
et al., 1962; Miles and Harms, 1982). The amount of phosphorus needed each day (Table 2-3) has been decreased from
amounts recommended in earlier editions. A daily intake of 250 mg of nonphytate phosphorus should be adequate for normal
production and health. Although feeding diets containing excess phosphorus is generally undesirable, poultry encountering
heat stress may require additional phosphorus. Garlich et al. (1978) and McCormick et al. (1980) reported that chickens fed
diets containing relatively high phosphorus levels were more tolerant of high ambient temperatures than were those fed
normal phosphorus levels. The use of dietary phosphorus at requirement levels should result in less phosphorus in excreta.
This fact may assume more importance in the future if manure application rates to land are determined on the basis of
phosphorus content.
Research information published about vitamin requirements does not indicate the need for any major change in
recommendations from the previous edition. However, results from several reports showed that, for maximum egg yield, the
choline requirement was about 1,050 mg per hen daily (Parsons and Leeper, 1984; Keshavarz and Austic, 1985; Miles et al.,
1986). Therefore the choline requirement for laying hens has been increased.
Brown-Egg-Laying Layers
Estimated nutrient requirements of brown-egg layers are listed in Table 2-3. Because little research has been
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done with brown-egg-laying layers, the committee had little quantitative information to review for establishing nutrient
requirements. Estimates of daily requirements given in Table 2-3 are listed as 10 percent greater than those of the white-egglaying layers. The 10 percent increase is justified on the basis that brown-egg-laying layers have heavier body weights and
generally produce more egg mass per hen daily.
Egg-Type Breeders
Nutrient requirements for egg-type breeders are listed in Table 2-3. Major nutrient requirements are the same for
producing an egg for human consumption as for producing an egg for hatching; however, dietary levels of trace minerals and
vitamins that result in maximum egg yield per day may be too low for the developing embryo (Naber, 1979). Vitamin and
trace mineral levels in the egg can be increased by increasing the dietary levels. Higher riboflavin, pantothenic acid, and
vitamin B12 levels are especially critical for maximum hatchability, although several other nutrients may also become
limiting. As a result, several of the micronutrient requirements are higher in breeding diets than in laying diets.
Molting Hens
After 8 to 12 months of egg production, some flocks are molted as a means of extending the period of production
(Zimmerman and Andrews, 1987). A combination of feed, water, and light restriction is usually used to stop egg production
and cause a rest, which may last from 3 to 6 weeks. A rest can also be induced by free-choice feeding of a diet containing a
deficiency or excess of a specific nutrient. Examples of nutrients used to induce molt include excess iodine (Arrington et al.,
1967), excess zinc (Supplee et al., 1961), and sodium chloride deficiency (Whitehead and Shannon, 1974; Naber et al., 1984).
After the rest, egg production can be initiated by stimulatory lighting. Little research information is available on the nutrient
requirements of molted hens; therefore the committee has assumed that requirements are similar to those of hens during the
first cycle of production.
TABLE 2-5 Typical Body Weights, Feed Requirements, and Energy Consumption of Broilers
Body Weight (g)
Weekly Feed
Cumulative Feed
Weekly Energy
Age
Consumption (g)
Consumption (g)
Consumption (kcal
(weeks)
ME/bird)
Male
Female
Male
Female
Male
Female
Male
Female
1
152
144
135
131
135
131
432
419
2
376
344
290
273
425
404
928
874
3
686
617
487
444
912
848
1,558
1,422
4
1,085
965
704
642
1,616
1,490
2,256
2,056
5
1,576
1,344
960
738
2,576
2,228
3,075
2,519
6
2,088
1,741
1,141
1,001
3,717
3,229
3,651
3,045
7
2,590
2,134
1,281
1,081
4,998
4,310
4,102
3,459
8
3,077
2,506
1,432
1,165
6,430
5,475
4,585
3,728
3,551
2,842
1,577
1,246
8,007
6,721
5,049
3,986
9
Cumulative Energy
Consumption (kcal
ME/bird)
Male
Female
432
419
1,360
1,293
2,918
2,715
5,174
4,771
8,249
7,290
11,900
10,335
16,002
13,794
20,587
17,522
25,636
21,508
NOTE: Values are typical for broilers fed well-balanced diets providing 3,200 kcal ME/kg.
MEAT-TYPE CHICKENS
Dietary requirements for meat-type chickens vary according to whether the birds are broilers being started and grown for
market, broiler breeder pullets and hens, or broiler breeder males.
Starting and Growing Market Broilers
Chickens of broiler strains have been selected for rapid weight gain and efficient utilization of feed. Broilers are usually
allowed to feed on an ad libitum basis to ensure rapid development to market size, although some interest has been expressed
in controlling feed intake in an attempt to minimize the development of excessive carcass fat. Broilers are marketed at a wide
range of ages and body weights (Table 2-5). Females may be grown to 900- to 1,000-g body weight to supply Cornish hens,
mixed sexes may be reared to 1.8 to 2 kg for use as whole birds and specialty parts, and males may be grown to 2.8 to 3 kg
for deboned meat. Thus it is difficult to establish a single set of requirements that is appropriate to all types of broiler
production. Furthermore, nutrient requirements may vary according to the criterion of adequacy. In the instance of essential
amino acids, greater dietary concentrations may be required to optimize efficiency of feed utilization than would be needed to
maximize weight gain. There also is evidence that the dietary requirement for lysine to maximize yields of breast meat of
broilers is greater than that needed to
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maximize weight gain (Acar et al., 1991) and that differences exist among strains of broilers with respect to this need for
more lysine (Bilgili et al., 1992).
Expression of a requirement for any nutrient is relative, and many factors must be considered. Many nutrients are
interdependent, and it is difficult to express requirements for one without consideration of the quantity of the other. Examples
include the relationships that exist between lysine and arginine and among calcium, phosphorus, and vitamin D3 levels in the
diet.
Other factors that may affect requirements include age and gender of the animal. Some studies suggest that males
require greater quantities of nutrients than do females at a similar age; however, when expressed as a percentage of the diet,
there seems to be little difference in nutrient requirements of the sexes. The requirements for many nutrients seem to diminish
with age, but for most nutrients there have been few research studies designed to precisely estimate requirements for all age
periods, especially for those beyond 3 weeks of age.
Any expression of nutrient requirements can be only a guideline representing a consensus of research reports. These
guidelines must be adjusted as necessary to fit the wide variety of ages, sexes, and strains of broiler chickens.
The values given in Table 2-6 are generally minimum levels that satisfy general productive activities and(or) prevent
deficiency syndromes. Requirements are presented for specific age periods. These age periods are based on the chronology
for which research data were available. These nutrient requirements are often implemented for younger age intervals or on a
weight-of-feed consumed basis. Where information is lacking, bold italicized values represent an estimate based on values
attained for other ages or related species. The data from the peer-reviewed scientific literature that serve as a basis for the
committee's estimation of nutrient requirements are presented in Appendix Table A-3a.
Amino Acids
Relatively high concentrations of dietary amino acids are needed to support the rapid growth of meat-type chickens.
Body weights of commercial meat-type chickens will increase 50- to 55-fold by 6 weeks after hatching. A large part of this
increase in weight is tissue of substantial protein content. Thus, adequate amino acid nutrition is vital to the successful
feeding program for this type of chicken.
Methionine plus Cystine
The greatest disagreement concerning amino acid requirements for broilers centers on the sulfur amino acids,
methionine and cystine. In
TABLE 2-6 Nutrient Requirements of Broilers as Percentages or Units per Kilogram of Diet (90 percent dry matter)
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part, this is because most studies are not designed to determine both the requirements of methionine per se and the
requirement for the combined quantity of methionine and cystine. Many attempts have been made, especially with purified
diets, to ascertain the relative proportions needed of these two amino acids, with variable results. Many have attributed a
share of the disagreement in estimated requirements to factors such as the sparing effects of choline (Quillen et al., 1961;
Pesti et al., 1979) or sulfate (Gordon and Sizer, 1955; Ross and Harms, 1970) or the negative effects of copper sulfate (Baker
and Robbins, 1979).
It is unfortunate that although a number of studies have been carried out to examine the effects of different dietary
variables on the requirement for methionine, few of these actually made attempts to estimate an overall requirement value.
Although calculations can be made in some instances, these do not have the statistical basis that values derived from the
original data would have had.
Another factor that may contribute to the disagreement in results is the comparison of results using crystalline amino
acid diets with results using diets based on practical ingredients, primarily corn and soybean meal. Although this difference
may relate in part to the incomplete digestion of the protein in the intact ingredients, most recent digestibility studies suggest
that amino acids in corn and soybean meal are well digested, on the order of 85 percent or more. Differences in digestibility
of practical and semipurified diets are, therefore, not of sufficient magnitude to account for the major differences that seem to
occur between these types of diets.
The cystine status of the basal diet is a major factor that contributes to the apparent disagreement in results, especially
when diets with intact ingredients are used. Generally, a basal diet, considered deficient in sulfur amino acids, is
supplemented with graded levels of methionine and the response determined. The point of maximum response is then noted,
and the sum of dietary plus supplemental methionine is added to the dietary cystine content to arrive at the need for total
sulfur amino acids (TSAA). However, this procedure assumes that the basal diet does not contain a surfeit of cystine.
Therefore one must determine whether or not the basal diet is adequate or excessive in cystine before combining these values
for a total TSAA estimate. Total dietary cystine levels can be influenced by dietary protein levels, choice of proteincontributing ingredients, and use of supplemental amino acids. Unfortunately, the majority of the reports estimate TSAA
requirements and do not attempt to differentiate between needs for methionine and needs for TSAA.
For methionine per se, there is minimal research on which to base changes in the recommendation of 0.5 percent made
in the previous edition. Of the reports in the literature for methionine requirements for the period from 0 to 21 days, two
(Waldroup et al., 1979; Tillman and Pesti, 1985) are above the NRC (1984) recommendation, four (Dean and Scott, 1965;
Robbins and Baker, 1980a; Moran, 1981; Thomas et al., 1985) are at or near that recommendation, and two (Klain et al.,
1960; Hewitt and Lewis, 1972) are considerably below. For the period of 3 to 6 or 6 to 8 weeks, there is even less work on
the requirements for methionine per se. The report of Moran (1981) plus estimates from a computer model (Hurwitz et al.,
1978) would support retaining the previously recommended value until sufficient research has been conducted to support its
modification.
Even greater diversity exists among estimates for TSAA requirements, as would be expected from the factors indicated
above. Evaluation of results obtained from feeding crystalline amino acid diets certainly suggests a markedly lower TSAA
value (Klain et al., 1960; Dean and Scott, 1965; Graber et al., 1971; Robbins and Baker, 1980a; Willis and Baker, 1980,
1981a; Baker et al., 1983). Although basing TSAA requirements on data using crystalline amino acids is perhaps not
justifiable for practical diets, it does point out that the TSAA requirement could be less if a proper balance between available
methionine and cystine existed.
In evaluating results from birds fed diets with intact ingredients, one can find values that support the change in
recommended TSAA requirements for 0 to 3 weeks of age from 0.93 to 0.87 percent of the diet (Nelson et al., 1960; Hewitt
and Lewis, 1972; Boomgaardt and Baker, 1973b,c; Woodham and Deans, 1975; Attia and Latshaw, 1979; Robbins and
Baker, 1980a,b; Wheeler and Latshaw, 1981; Baker et al., 1983; Mitchell and Robbins, 1983; Thomas et al., 1985). In many
of these studies, diets were supplemented with lysine, which permitted a lower protein level and reduced cystine content;
therefore a surfeit of cystine was less likely to exist in these studies. Research is needed using practical ingredients to
evaluate the separate needs for methionine and cystine in such diets.
For the 3- to 6-week period, most reports are in agreement with the previous recommendation (Graber et al., 1971;
Holsheimer, 1981; Wheeler and Latshaw, 1981; Mitchell and Robbins, 1983). Two reports (Jensen et al., 1989; Mendonca
and Jensen, 1989a) suggested a higher value, based in part on reduction in carcass fat content. There is minimal research on
the TSAA needs from 6 to 8 weeks of age and little justification for change in the previous recommendation. More research is
needed to delineate the separate needs for methionine and cystine in diets consisting of practical ingredients. This research
may eliminate much of the current disagreement regarding TSAA needs of the broiler.
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Arginine
The committee has made significant changes in its recommendation for the arginine requirements of broilers. It has
eliminated from consideration all studies in which potential lysine:arginine antagonisms existed because such antagonisms
are unlikely to occur with practical ingredients. Recommended requirements have been reduced to 1.25 and 1.1 percent for
the 0-to 3- and 3- to 6- week growth periods, respectively.
Lysine
The requirement of broilers from 0 to 3 weeks of age has been reduced from 1.2 to 1.1 percent of the diet. There has
been little recent research on the requirements for this amino acid, but evaluation of previous research supports this reduction
(Edwards et al., 1956; Boomgaardt and Baker, 1973a,b; Woodham and Deans, 1975; McNaughton et al., 1978; Burton and
Waldroup, 1979). There is a dearth of published recommendations for the period from 3 to 6 weeks of age. Limited research,
however, supports the previous recommendation (Holsheimer, 1981). Research results for the period from 6 to 8 weeks are
inconclusive. Some work suggests that the previous requirement is low (Bornstein, 1970; Boomgaardt and Baker, 1973b),
whereas other studies suggest that it is high (Chung et al., 1973; Twining et al., 1973; Thomas et al., 1977). Therefore, the
previous requirement of 0.85 percent was not changed.
Tryptophan
The committee has reduced the requirement for this amino acid from 0.23 to 0.2 percent for the broiler 0 to 3 weeks of
age on the basis of its evaluation of published reports from many sources (Wilkening et al., 1947; Griminger et al., 1956;
Klain et al., 1960; Boomgaardt and Baker, 1971; Hewitt and Lewis, 1972; Woodham and Deans, 1975; Steinhart and
Kirchgessner, 1984; Smith and Waldroup, 1988a). Minimal research has been conducted on tryptophan requirements of the
broiler at more than 3 weeks. Estimates from computer modeling (Hurwitz et al., 1978) suggest that lower levels of
tryptophan may be required during this period, but these estimates have not been rigorously examined.
Threonine
Considerable work has been conducted on the threonine requirement for broiler chickens in recent years. The majority of
the studies support the present recommended value of 0.8 percent for broilers at 0 to 3 weeks of age (Uzu, 1986; Robbins,
1987; Thomas et al., 1987; Bertram et al., 1988; Smith and Waldroup, 1988b; Austic and Rangel-Lugo, 1989). Little research
has been done on threonine requirements for broilers older than 3 weeks of age.
Isoleucine, Leucine, Valine, Phenylalanine, Phenylalanine plus Tyrosine, Glycine plus Serine, Histidine, and Proline
Sufficient studies with intact protein diets have been conducted to allow estimation of the requirements for leucine,
isoleucine, and valine during the 0-to 3-week period (Almquist, 1947; D'Mello, 1974; Woodham and Deans, 1975; Thomas et
al., 1988). Only a few studies with intact protein diets have been conducted for phenylalanine or phenylalanine plus tyrosine
(Almquist, 1947; Woodham and Deans, 1975) and for glycine plus serine (Ngo and Coon, 1976) during the period from 0 to
3 weeks. Therefore the committee considered studies with purified diets (Fisher et al., 1957; Klain et al., 1960; Dean and
Scott, 1965; Sasse and Baker, 1972; Coon et al., 1974; Baker et al., 1979) in estimating these requirements. The reported
values for phenylalanine plus tyrosine and glycine plus serine vary greatly among studies, particularly in the latter instance.
The histidine requirement for the period from 0 to 3 weeks is based primarily on purified diet studies (Klain et al., 1960;
Dean and Scott, 1965; Baker et al., 1979). Although proline is not usually considered to be an essential amino acid for
poultry, research has shown that young chicks may not synthesize sufficient proline to meet their requirements (Greene et al.,
1962; Graber et al., 1970); thus, a dietary source of proline must be provided.
The committee found no published research data for this group of amino acids for the periods from 3 to 6 and 6 to 8
weeks, although the study by Mendonca and Jensen (1989b) suggested that the valine requirement for 3 to 6 weeks exceeds
0.70 percent. Since the lysine requirements for these growth periods are documented, the requirements for this group of
amino acids for the periods from 3 to 6 and 6 to 8 weeks have been estimated from the lysine values by using the amino
acid:lysine ratio for the period from 0 to 3 weeks. Thus the committee assumed that the ratios or patterns between these
amino acids and lysine are relatively consistent throughout the growth stages.
Minerals
The extent of research conducted on different minerals and vitamins is often in direct proportion to their economic value
or to the likelihood of encountering a dietary deficiency in practical diets. Thus there is a great deal of literature concerning
the calcium and phosphorus requirements of the broiler and minimal research concerning requirements for trace elements.
The precise requirements for minerals such as potassium, magnesium, and iron in practical diets are not well defined because
practical diets are usually adequate or only slightly deficient in these minerals. The requirements for minerals such as iron,
manganese, and zinc are much lower for chicks fed semipurified diets containing little or no phytate and fiber than for those fed
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practical diets, mainly because of relatively poor bioavailability of some minerals in practical ingredients (Kratzer and Vohra,
1986). For example, the bioavailability of manganese is very low in most practical feedstuffs, and there is evidence that
practical ingredients reduce the bioavailability of inorganic dietary manganese (Halpin and Baker, 1986). The bioavailability
of minerals in inorganic mineral supplements also varies greatly. For example, the bioavailability of zinc in zinc sulfate is
much higher than in zinc oxide (Wedekind and Baker, 1990). Consequently, the reported requirement for a mineral may vary
among studies owing to differences in the bioavailability of the supplemental mineral source and the use of ingredients that
interfere with utilization of the mineral under study.
Although substantial research has been conducted for most vitamins, the requirements for practical diets are not well
defined. Practical diets are not markedly deficient in some vitamins. Consequently, several of the vitamin requirements are
extrapolated from studies with purified or semipurified diets. The dietary levels needed to maximize some parameters may be
higher than those needed to maximize growth. Examples of the latter include vitamin D3 levels for maximum tibia ash
(Waldroup et al., 1963a; Lofton and Soares, 1986), vitamin E levels for maximum immune response (Tengerdy and Nockels,
1973; Colnago et al., 1984), and riboflavin levels for prevention of leg paralysis (Ruiz and Harms, 1988a). It is generally
assumed that vitamin requirements decrease with increasing age, although this relationship is not well documented with the
exception of choline in purified diets.
Calcium and Phosphorus
No changes have been made in the previously recommended calcium requirement of the broiler chick. Requirements for
phosphorus are expressed in terms of nonphytate phosphorus. The nonphytate phosphorus requirement for the chick at 0 to 3
weeks of age remains unchanged; however, recommended values for 3 to 6 and 6 to 8 weeks have been reduced on the basis
of studies by O'Rourke et al. (1952), Waldroup et al. (1963b, 1974a), Twining et al. (1965), Sauveur (1978), Yoshida and
Hoshii (1982a), and Tortuero and Diez Tardon (1983).
Potassium, Sodium, and Chlorine
A reduction has been made in the potassium requirement of the broiler. The potassium requirement of broilers fed a
semipurified diet seems to be between 0.25 and 0.30 percent (Leach et al., 1959). The requirement for broilers fed a practical
diet is not documented. The requirements for sodium and chlorine have been increased for the period from 0 to 3 weeks on
the basis of recent studies. The requirements for these minerals seem to decrease with increasing age (Hurwitz et al., 1973;
Edwards, 1984). The research of Edwards (1984) has justified a reduction in the levels of sodium and chlorine recommended
for broilers at 6 to 8 weeks of age.
Magnesium
The reported requirement varies among studies. Part of this variation may be due to the calcium and phosphorus content
of the diet. Although type of diet varies among studies, there does not seem to be a consistent relationship between diet type
and the reported magnesium requirement. After 3 weeks of age, the values suggested by the committee are only estimates.
Iron and Copper
Although only a few studies have been conducted on iron requirements of broilers, the results are consistent and indicate
that the requirement is approximately 80 mg/kg (Davis et al., 1968; McNaughton and Day, 1979). Southern and Baker (1982)
report that the requirement was only 40 mg/kg for chicks fed a dextrose-casein diet. The copper requirement of 8 mg/kg is
based on the study of McNaughton and Day (1979). The committee suggests only estimated values after 3 weeks of age.
Manganese
Values given for chicks of all ages show wide differences in requirements depending on the type of diet used. The
requirement reported for chicks fed a semipurified dextrose-casein diet (14 mg/kg; Southern and Baker, 1983a) is much lower
than that of chicks fed a diet containing practical ingredients (50 mg/kg/ Gallup and Norris, 1939a,b).
Zinc
The zinc requirement of the young broiler is approximately 35 to 40 mg/kg in semipurified diets containing isolated soy
protein or casein (Morrison and Sarett, 1958; O'Dell et al., 1958; Roberson and Shaible, 1958). Studies on corn-soybean meal
and sesame meal diets suggest that the requirement is in excess of 40 mg/kg (Edwards et al., 1959; Lease et al., 1960; Zeigler
et al., 1961). This conclusion was based primarily on small growth responses to zinc supplementation of the basal diets. The
estimated zinc requirement is somewhat tenuous, because the estimate was based on calculated values for zinc content of the
feed ingredients. Recent work by Wedekind et al. (1990) showed that the tibia zinc concentration of chicks fed a cornsoybean meal diet was increased markedly by dietary zinc supplementation but did not provide an estimate of requirements.
The source of supplemental zinc used in most of the cited studies was zinc sulfate or zinc chloride. Availability of zinc varies
among sources (Wedekind and Baker, 1990). In a diet containing egg white as the primary protein source, the requirement for
zinc is only 14 to 18 mg/kg (Southern and Baker, 1983b; Dewar and Downie, 1984). Only tentative values are given for
chicks after 3 weeks of age.
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Iodine
Little research has been conducted to establish the iodine requirement of the broiler chick. The present requirement is
based on the study by Creek et al. (1957).
Selenium
No changes have been made in the recommended dietary selenium concentrations for broiler chickens. A concentration
of 0.15 mg selenium per kilogram of diet is recommended (Jensen et al., 1986).
Vitamins
Vitamin A
Tentative requirement values have been listed for all ages. The requirement estimates vary from 900 to 2,200 IU/kg
among studies. Requirement values from more recent studies are lower than those from earlier ones.
Vitamin D
The requirement estimates for maximum growth are consistent among most studies. The requirement for maximum tibia
ash, however, may be higher than that for growth (Waldroup et al., 1965; Lofton and Soares, 1986).
Vitamin E
Tentative values have been expressed for all ages. The results of the few studies conducted are variable. The requirement
for prevention of encephalomalacia may be higher than that for growth only (Singsen et al., 1955). In addition, the
requirement for maximum immune response may be much higher than that for growth (Tengerdy and Nockels, 1973;
Colnago et al., 1984).
Vitamin K
The vitamin K requirements of the broiler are unchanged. The requirement is estimated at approximately 0.5 mg/kg for
chicks fed glucose-isolated soy protein diets (Nelson and Norris, 1960, 1961b).
Riboflavin
The riboflavin requirements for broilers at 0 to 3 and 3 to 6 weeks of age (3.6 mg/kg of diet) are unchanged. Most
studies indicate that the riboflavin requirement is 2.5 to 3.5 mg/kg. Several studies have indicated that the requirement for
prevention of leg paralysis is higher than that for growth (Ruiz and Harms, 1988c).
Pantothenic Acid
Tentative requirements have been expressed for broilers of all ages. Little work has been done, and there is no good
basis for the requirement in practical diets. The requirement is 5 mg/kg in a purified diet, and thus twice this level should be
adequate for practical diets to compensate for potentially limited availability of pantothenic acid from the ingredients.
Bauernfeind et al. (1942) reported that 7.5 to 10 mg of pantothenic acid per kilogram of diet was adequate for Leghorn chicks
and that practical diets normally contain sufficient levels of this vitamin. Jukes and McElroy (1943) also reported a
pantothenic acid requirement of 10 mg/kg of diet.
Niacin
The niacin requirement has been increased for broilers of all ages (see Table 2-5). Requirement estimates vary from 22
to greater than 55 mg/kg among studies using intact protein diets, with most estimates being in the range of approximately 25
to 35 mg/kg. The requirement is somewhat lower for purified diets (Ruiz and Harms, 1988a; 1990).
Vitamin B12
Few requirement studies have been conducted. The requirement seems to be approximately 0.01 mg/kg (Looi and
Renner, 1974; Rys and Koreleski, 1974).
Choline
No changes have been made in the choline requirement of the broiler at 0 to 3 weeks of age, and tentative requirements
are given for broilers at 3 to 6 and 6 to 8 weeks. Many studies have been conducted on choline requirements, and the
requirement estimates are highly variable. Choline requirements are influenced by protein and sulfur amino acid content of
the diet and by age of broilers. The requirements listed in Table 2-5 should be sufficient for practical diets containing
adequate levels of methionine and cystine. The choline requirement is much lower and decreases markedly with increasing
age for chicks fed purified diets (Molitoris and Baker, 1976; Lowry et al., 1987). A decrease in choline requirement with age
has not been documented when practical diets are fed. Requirement values for broilers from 3 to 6 and 6 to 8 weeks,
however, have been extrapolated from studies that used purified diets (Gardiner and Dewar, 1976; Molitoris and Baker, 1976;
Lowry et al., 1987).
Biotin
No changes have been made in the biotin requirement of the broiler to 6 weeks of age, with a tentative requirement
expressed for 6 to 8 weeks. Estimates from most studies indicate that the requirement is between 0.15 and 0.20 mg/kg.
Folic Acid
No changes have been made in the folic acid requirement of the broiler at 0 to 3 and 3 to 6 weeks of age, with tentative
requirements expressed for 6 to 8 weeks. Requirement values vary among studies. Recent studies, however, indicate that the
requirement is between 0.35 and 0.50 mg/kg when determined with semipurified diets. Thus the requirement is probably
higher when birds are fed practical diets.
Thiamin
Tentative requirements are expressed for broilers of all ages. There is little research with broilers on which to base a
requirement. The requirement seems to
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be relatively low, and practical diets normally contain levels well in excess of the estimated requirements.
Pyridoxine
The pyridoxine requirement has been increased for broilers of all ages, with a tentative requirement given for broilers at
6 to 8 weeks of age. Many studies have been conducted, with requirement estimates ranging from 2.3 to 3.5 mg/kg for intact
protein diets. The requirement seems to be only approximately 1.0 mg/kg for a purified diet (Lee et al., 1976; Yen et al.,
1976). The pyridoxine requirement, however, increases with an increase in dietary protein level (Gries and Scott, 1972a;
Daghir and Shah, 1973).
Essential Fatty Acid
Linoleic Acid
The linoleic acid requirement has been estimated as 1.0 percent of the diet (Balnave, 1970).
Broiler Breeder Pullets and Hens
Meat-type breeder hens will become obese if allowed ad libitum consumption of feed; therefore some form of nutrient
limitation must be practiced. Most research has focused on feeding systems, with some form of quantitative restriction of
intake generally practiced to maintain body weights within guidelines suggested by the breeder. Early research suggested that
feeding bulky, high-fiber diets would successfully limit MEn intake (Milby and Sherwood, 1953; Singsen et al., 1959; Isaacks
et al., 1960; Summers et al., 1967; Fuller et al., 1973), but more recent studies indicate that modern broiler strains can
consume large volumes of feed, a capability that makes this method impractical as a means of controlling weight (Waldroup
et al., 1976a). Other studies have suggested that low-protein diets (Waldroup et al., 1966), diets low in specific amino acids
(Singsen et al., 1964), or diets imbalanced in amino acids (Couch and Abbott, 1974) might control body weight when offered
for ad libitum consumption, but such diets have not been readily accepted in commercial practice because of large variability
in bird response.
Little research has been conducted to determine the specific nutrient requirements of meat-type females from hatch to
maturity. Powell and Gehle (1975) estimated the tryptophan requirement of growing broiler breeder pullets; this seems to be
the lone estimate of protein or amino acid needs during this age period. Harms (1980) and Harms and Wilson (1987) have
suggested requirements for the growing pullet, but these have not been subjected to rigid evaluation. Therefore there is not
sufficient research data on which to base suggested requirements for the growing and developing broiler breeder meat-type
pullet at this time.
Nutrient requirement data presented in Table 2-7 for the broiler breeder meat-type hen are limited to those for which
some documentation is available.
Protein And Amino Acids
Chickens do not require a specific level of crude protein per se; rather, they have a requirement for specific amino acids
plus sufficient protein to supply either the nonessential amino acids themselves or amino nitrogen for their synthesis. In the
instance of meat-type breeder hens, there is a paucity of research directed toward determining specific requirements for
essential amino acids. Therefore a minimum crude protein intake is generally designated to provide adequate amounts of
essential amino acids whose requirements are not adequately known.
Daily crude protein intakes of 18 to 20 g per hen seem adequate, assuming that essential amino acid needs are met
(Waldroup et al., 1976b; Pearson and Herron, 1981; Spratt and Leeson, 1987), although more abundant levels (up to 23 g/
day) may be needed during periods of highest productivity to achieve maximum egg mass yield (Jeroch et al., 1982; Schloffel
et al., 1988). Because the size of the
TABLE 2-7 Nutrient Requirements of Meat-Type Hens for Breeding Purposes as Units per Hen per Day (90 percent dry matter)
Unit
Requirements
Nutrient
Protein and amino acids
a
g
19.5
Protein
Arginine
mg
1,110
Histidine
mg
205
Isoleucine
mg
850
Leucine
mg
1,250
Lysine
mg
765
Methionine
mg
450
Methionine + cystine
mg
700
Phenylalanine
mg
610
Phenylalanine + tyrosine
mg
1,112
Threonine
mg
720
Tryptophan
mg
190
Valine
mg
750
Minerals
Calcium
g
4.0
Chloride
mg
185
Nonphytate phosphorus
mg
350
Sodium
mg
150
Vitamin
µg
16
Biotin
NOTE: These are requirements for hens at peak production. Broiler breeder hens are usually fed on a controlled basis to maintain body
weight within breeder guidelines. Daily energy consumption varies with age, stage of production, and environmental temperature but
usually ranges between 400 and 450 ME kcal per hen at peak production. For nutrients not listed, see requirements for egg-type breeders
(Table 2-3) as a guide. Where experimental data are lacking, values typeset in bold italics represent an estimate based on values obtained
for other ages or related species.
a
Broilers do not have a requirement for crude protein per se. There, however, should be sufficient crude protein to ensure an adequate
nitrogen supply for synthesis of nonessential amino acids. Suggested requirements for crude protein are typical of those derived with
corn-soybean meal diets, and levels can be reduced somewhat when synthetic amino acids are used.
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33
egg has a significant effect on the initial weight of the chick and its subsequent performance (Gardiner, 1973; Guill and
Washburn, 1973; Proudfoot and Hulan, 1981), maximum egg weight during early production is an important economic
factor. The protein requirement for dwarf breeder hens does not exceed 13.6 percent of the diet (Larbier et al., 1979).
Excessive crude protein intakes are to be avoided. Daily intakes of 27 g per hen had adverse effects on hatchability
(Pearson and Herron, 1981, 1982). Lower crude protein intakes may be satisfactory if additional amino acid supplementation
is practiced. Bornstein et al. (1979) calculated that a daily crude protein intake of 15.6 to 16.5 g per hen would be sufficient in
terms of an ideal amino acid mixture. Performance of hens fed corn-soybean meal diets providing 16 g protein per day was
not improved by supplemental lysine and methionine (Waldroup et al., 1976b).
Few trials have been conducted to determine specific amino acid requirements. Harms and Wilson (1980) reported a
daily requirement for methionine of between 400 and 478 mg; 400 mg per day gave performance statistically equivalent to
that at higher levels of intake. Halle et al. (1984), using nitrogen balance studies, indicated a TSAA need of 694 mg per day.
For dwarf (dw) hens, Guillaume (1977) estimated daily methionine and lysine needs of 360 to 380 and 750 mg per hen,
respectively.
Wilson and Harms (1984) obtained satisfactory performance with average daily intakes per hen of 682 mg of TSAA,
808 mg of lysine, 1,226 mg of arginine, and 223 mg of tryptophan, with 18.6 g of crude protein per day. Using various
prediction models or equations, several workers have estimated amino acid requirements (Waldroup and Hazen, 1976;
Waldroup et al., 1976c; Scott, 1977; Bornstein et al., 1979). In the study by Bornstein et al. (1979), hens fed diets formulated
to meet these requirements on the basis of prediction models performed as well as those fed diets formulated in the
conventional way.
Energy
Broiler breeder hens are usually fed on a controlled basis to maintain body weight within breeder guidelines. Daily
energy consumption will vary with age, stage of production, and environmental temperature, but will usually range from 400
to 450 kcal ME per hen daily (Waldroup and Hazen, 1976; Waldroup et al., 1976a; Bornstein et al., 1979; Bornstein and Lev,
1982; Pearson and Herron, 1982; Spratt and Leeson, 1987; Spratt et al., 1990a,b).
Minerals And Vitamins
Calcium
Shell strength of eggs from meat-type hens increases as calcium level is increased (Mehring, 1965). Egg production and
hatchability of meat-type hens on litter were not improved by feeding more than 3.91 g of calcium per hen daily (Wilson et
al., 1980). One of the best determinants of calcium adequacy for breeder hens is egg specific gravity; eggs should have a
specific gravity of 1.080 or greater for optimal hatchability (McDaniel et al., 1979). Since meat-type hens are usually given a
daily allotment of feed early in the morning before significant eggshell calcification occurs, supplying a portion of the
calcium in an afternoon feeding may improve eggshell quality (Farmer et al., 1983; Van Wambeke and DeGroote, 1986).
Feeding the entire dietary allocation in the afternoon, however, may significantly reduce hatchability because of production
of eggs with thicker eggshells (Brake, 1988).
Phosphorus
No significant differences in egg production, hatchability of fertile eggs, or specific gravity of eggs were noted in
feeding from 532 to 1,244 mg total phosphorus per hen daily (163 to 863 mg nonphytate phosphorus per hen daily), although
egg production was improved numerically by feeding 718 mg total phosphorus (338 mg nonphytate phosphorus) per day
(Wilson et al., 1980). For both calcium and phosphorus, requirements for hens maintained in cages may be significantly
greater than for hens on litter floors (Harms et al., 1961; Singsen et al., 1962; Harms et al., 1984).
Sodium
Egg production, feed efficiency, egg weight, fertility, and hatchability of meat-type breeder hens were not improved by
feeding more than 154 mg of sodium per hen daily (Damron et al., 1983); sodium intakes in excess of 320 mg per day were
shown to reduce fertility.
Chlorine
Harms and Wilson (1984) reported that 254 mg of chlorine per hen daily resulted in the best overall performance of
meat-type broiler hens, as measured by egg production and hatchability. However, performance on this intake did not differ
significantly from performance on intakes of 185 mg per day.
Biotin
The requirement for biotin by the meat-type hen has been estimated to be 16 µg per hen daily. The hen may be
considered to be receiving adequate biotin if the yolk biotin concentration is at least 550 ng/g (Whitehead et al., 1985).
Broiler Breeder Males
Historically, meat-type breeder cockerels have been grown with the females. Because of recent changes in genetics and
management practices, an increasing number of males are being grown or fed separately. Males maintained in floor pens with
natural mating may be fed from a separate feeding system; males maintained in cages for artificial
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NUTRIENT REQUIREMENTS OF CHICKENS
34
insemination may be individually fed. The major advantage of separate feeding is control of body weight and its subsequent
impact on fertility and mating ability. Thus a set of nutrient requirements for male meat-type breeders, although limited in
scope, is listed in Table 2-8. It should be noted that diets intended for use by the breeder hen, when fed to control male body
weight, appear to have no detrimental effects on male performance.
Protein
Protein requirements of breeder cockerels have been evaluated during the growing and adult periods by using both
White Leghorn and Meat-type cockerels. In studies with Single Comb White Leghorn (SCWL) cockerels, low crude protein
levels fed during the grower period reduced body weights and delayed testicular development, but, on subsequent feeding of
adequate protein, reproductive performance was not impaired (Wilson et al., 1965; Jones et al., 1967). Diets containing 12.4
percent crude protein offered for ad libitum consumption to broiler breeder males during the period of 7 to 21 weeks of age
were adequate for development of the reproductive system and subsequent reproductive performance (Wilson et al., 1971).
Broiler breeder males can be fed 12 to 14 percent crude protein on a restricted basis after 4 weeks of age with no adverse
effects on final body weight, sexual maturity, or semen quality; a greater number of males produced semen through 53 weeks
when fed 12 percent crude protein than when fed higher levels (Wilson et al., 1987a). In a subsequent study (Wilson et al.,
1987b), a 9 percent crude protein diet fed beginning at 43 days and continuing through 50 weeks was adequate to support
maximum reproductive performance. In both these studies, amino acid content was maintained at a constant percentage of the
protein level. There were no differences in semen characteristics of broiler breeder males fed 12 to 18 percent crude protein
during the period from 4 to 20 weeks; males fed 15 percent crude protein during the period from 1 to 4 weeks had
significantly higher fertility from 24 to 27 weeks than did males fed 20 percent crude protein (Vaughters et al., 1987). Semen
production of broiler breeder males kept in cages can be maintained from 20 to 60 weeks on a daily protein intake of 10.9 to
14.8 g per day (Buckner and Savage, 1986).
TABLE 2-8 Nutrient Requirements of Meat-Type Males for Breeding Purposes as Percentages or Units per Rooster per Day (90 percent
dry matter)
Age (weeks)
Unit
0 to 4
4 to 20
20 to 60
Metabolizable energya
kcal
—
—
350 to 400
Protein and amino acids
%
15.00
12.00
—
Proteinb
%
0.79
0.64
—
Lysinec
c
%
0.36
0.31
—
Methionine
Methionine + cystinec
%
0.61
0.49
—
Minerals
Calcium
%
0.90
0.90
—
Nonphytate phosphorus
%
0.45
0.45
—
Protein and amino acids
Protein
g
—
—
12
mg
—
—
680
Argininec
c
mg
—
—
475
Lysine
mg
—
—
340
Methioninec
c
mg
—
—
490
Methionine + cystine
Minerals
Calcium
mg
—
—
200
mg
—
—
110
Nonphytate phosphorus
NOTE: For nutrients not listed, see requirements for egg-type pullets (Table 2-3) as a guide. Where experimental data are lacking, values
typeset in bold italics represent an estimate based on values obtained for other ages or related species.
a Energy needs are influenced by the environment and the housing system. These factors must be adjusted as required to maintain the
body weight recommended by the breeder.
b Broilers do not have a requirement for crude protein per se. There, however, should be sufficient crude protein to ensure an adequate
nitrogen supply for synthesis of nonessential amino acids. Suggested requirements for crude protein are typical of those derived with
corn-soybean meal diets, and levels can be reduced somewhat when synthetic amino acids are used.
c Amino acid requirements estimated by using the model of Smith (1978).
Energy
Daily energy intakes of 400 (McCartney and Brown, 1980) and 458 kcal ME per bird (Brown and McCartney, 1983)
have been reported as adequate for broiler breeder males maintained on litter. For broiler breeder males maintained in cages,
346 (Brown and McCartney, 1986) or 358 kcal ME per bird daily (Buckner et al., 1986) were sufficient.
Minerals
The calcium requirement of the breeder cockerel is much lower than that of the hen, but levels fed to the hen apparently
are not detrimental to the reproductive performance of the male. Wilson et al. (1969) indicated that the calcium requirement
of SCWL cockerels did not exceed 0.2 percent, but that levels as high as 3 percent were not detrimental. In calcium balance
studies with SCWL cockerels, Norris et al. (1972) found that the daily requirement was 7.98 mg per kg of body weight.
Kappleman et al. (1982) concluded that there were no differences in the reproductive performance of broiler breeder
cockerels fed 0.5 to 7 g of calcium daily per bird.
Phosphorus
Norris et al. (1972) found that diets containing 0.1 percent nonphytate phosphorus were satisfactory for SCWL
cockerels. Bootwalla and Harms (1989) found that no more than 110 mg of nonphytate phosphorus per bird daily were
needed for maintaining reproductive capacity and bone integrity in broiler breeder cockerels.
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NUTRIENT REQUIREMENTS OF TURKEYS
35
3
Nutrient Requirements of Turkeys
The nutrient requirements of turkeys are divided into needs of birds used as a source of growth and needs of those for
reproduction. These two categories differ largely in the proportion of nutrients devoted to productive use as opposed to those
used for maintenance activities.
Requirement values given in Table 3-1 are usually minimum levels that satisfy general productive activities and(or)
prevent deficiency symptoms. The values given often represent an approximation of values from more than one study. Where
information is lacking, italicized values represent an estimate based on values obtained for other ages or related species.
Values selected by the committee as best representing the requirement were those for which the research was recent and
performed under practical terms in which all nutrient needs in addition to the nutrient in question were satisfied. The
experimental data from the peer-reviewed scientific literature that are the basis for the committee's nutrient requirement
recommendations are given in Appendix Table A-4.
STARTING AND GROWING TURKEYS
The growth rate of turkeys has increased greatly during the past decade. Approximate live body weights per age and
feed consumption data of contemporary turkeys are shown in Table 3-2. Increased growth rates have occurred through the
efforts of the major commercial breeders, and parent stock has increased in size as well, particularly the hen. Further
processing of the carcass into convenience products also has expanded and now occupies the greatest part of total production.
Substantial improvements in the rates of gain and feed efficiencies of commercially available strains have occurred
during the last decade. The nutrient requirements given in Table 3-1 are based on earlier research and the chronological age
of the experimental turkeys used at that time. For the most part, these nutrient levels are still being employed by the industry
at large; however, because of improvements in growth rates these levels are now being used at earlier ages. Such changes
have not been experimentally verified as being appropriate, but commercial results indicate satisfactory performance.
Examples of these age adjustments for male and female turkeys are shown in Table 3-1, footnotes a and b, respectively.
Commercially available strains of turkey may differ in the chronology of their development. The nutrient requirements
given on Table 3-1 represent the approximate needs for development of large-type turkeys. Medium- and small-type turkeys
finish progressively earlier than the large. For the given nutrient levels to be employed effectively, those levels representing
each age interval should be provided according to the corresponding stages of development.
The requirements are expressed as concentrations in the feed. These concentrations are such that adequate total intake is
ensured and the nutrient balance is favorable. Both factors are necessary. A balanced feed having lower nutrient
concentrations than shown may not permit sufficient intake to meet the bird's absolute need. Conversely, an increased
concentration of nutrients ensures adequacy but may not be cost effective.
Pelleting is widely practiced in feed manufacturing, and feeding a pelleted diet usually leads to an improvement in
performance. Pelleting may increase nutrient digestibility in some constituent feedstuffs; however, the primary result is
improved use of the nutrients already available apparently because of reduced physical activity by the bird. Generally,
pelleting facilitates feed intake, increases net energy of production from metabolizable energy (ME), and reduces overall feed
wastage (Moran, 1989b). These benefits are accentuated as feed nutrient level decreases and as birds become progressively
older, provided the feed remains in pelleted form.
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36
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NUTRIENT REQUIREMENTS OF TURKEYS
37
TABLE 3-2 Growth Rate and Feed and Energy Consumption of Large-Type Turkeys
Age (weeks)
Body Weight (kg)
Feed Consumption per
Cumulative Feed
Week (kg)
Consumption (kg)
Male
Female
Male
Female
Male
Female
1
0.12
0.12
0.10
0.10
0.10
0.10
2
0.25
0.24
0.19
0.18
0.29
0.28
3
0.50
0.46
0.37
0.34
0.66
0.62
4
1.0
0.9
0.70
0.59
1.36
1.21
5
1.6
1.4
0.85
0.64
2.21
1.85
6
2.2
1.8
1.10
0.80
3.31
2.65
7
3.1
2.3
1.40
0.98
4.71
3.63
8
4.0
3.0
1.73
1.21
6.44
4.84
9
5.0
3.7
2.00
1.42
8.44
6.26
10
6.0
4.4
2.34
1.70
10.78
7.96
11
7.1
5.2
2.67
1.98
13.45
9.94
12
8.2
6.0
2.99
2.18
16.44
12.12
13
9.3
6.8
3.20
2.44
19.64
14.56
14
10.5
7.5
3.47
2.69
23.11
17.25
15
11.5
8.3
3.73
2.81
26.84
20.06
16
12.6
8.9
3.97
3.00
30.81
23.06
17
13.5
9.6
4.08
3.14
34.89
26.20
18
14.4
10.2
4.30
3.18
39.19
29.38
19
15.2
10.9
4.52
3.31
43.71
32.69
20
16.1
11.5
4.74
3.40
48.45
36.09
a
a
a
21
17.0
4.81
53.26
a
a
a
22
17.9
5.00
58.26
a
a
a
23
18.6
5.15
63.41
a
a
a
24
19.4
5.28
68.69
a
ME Consumption per Week
(Mcal)
Male
Female
0.28
0.28
0.53
0.5
1.0
1.0
2.0
1.7
2.5
1.9
3.2
2.3
4.1
2.8
5.0
3.5
6.0
4.3
7.0
5.1
8.0
5.9
9.0
6.8
9.9
7.6
10.8
8.4
11.6
9.0
12.3
9.6
13.1
10.1
13.8
10.5
14.5
10.9
15.2
11.2
a
15.9
a
16.5
a
17.1
a
17.4
No data given because females are usually not marketed after 20 weeks of age.
Energy
In calculating the total metabolizable energy for the complete feed, the metabolizable energies provided by each
feedstuff are assumed to be additive. The MEn content of the complete feed influences feed intake, which, in turn, may
influence the concentrations of most other nutrients that are needed to satisfy requirements. An inverse relationship exists
between the MEn concentration of the diet and feed consumption of turkeys. However, as discussed in Chapter 1 (Setting
Dietary Levels), changes in dietary MEn concentration and thus, the use of specific nutrient-to-dietary MEn ratios in
formulating turkey diets is questionable, especially when economical growth and feed efficiency are primary objectives (Pesti
and Fletcher, 1983; Sell et al., 1985; 1989).
The MEn levels given in Table 3-2 at each age period are not intended to be absolute but to establish a feed intake
reference for other nutrients. The energy and amino acid levels given would be satisfied largely when corn and soybean meal
are combined with a small amount of added fat, in turn permitting near-maximum growth. Nutrient levels may be increased
without adversely affecting performance; however, a moderate reduction in nutrient levels would likely require pelleting of
the associated feed to prevent adverse effects on growth rate.
Net energy of production is difficult to estimate because maintenance expenditures vary extensively. Environmental
temperature is one of the most influential factors affecting maintenance, which, in turn, may lead to changes in feed intake.
Changes in the maintenance energy requirement in response to environmental temperature may not be linear. Hurwitz et
al. (1980) observed that the maintenance energy requirement for both sexes of turkeys, during the period from 32 to 60 days
of age, was between 2.45 and 2.70 kcal/g67 of body weight at 12°C. This requirement progressively decreased from 12° to 24°
C, then remained constant between 24° and 28°C and increased thereafter through 35°C. The maintenance energy need in
response to temperature also differs with age. In a study on the 20-week-old male turkey, Hurwitz et al. (1983b) found the
requirement at 10°C to approximate 2.15 kcal/g67, but unlike the requirement for the younger bird (32 to 60 days) there was
an uninterrupted decrease through to 35°C. In both of these studies the advantage to net energy of production increased as
temperature increased; however, feed intake and growth were not altered accordingly.
Protein And Amino Acids
A protein requirement of 28 percent for starting poults is supported by the work of Lloyd et al. (1949), Atkinson et al.
(1957), Herz et al. (1975a), and Richter et al. (1980). Reduced levels of protein can decrease early growth, but if the protein
reduction is moderate, compensatory gain of large-type turkeys prior to marketing may overcome the deficit. The progressive
reduction in the protein requirement as the turkey grows is well established. A level of 12 percent protein with 2,900 kcal ME/
kg for holding turkeys prior to reproduction is consistent in terms of the protein:energy ratio with the 14 percent protein at
3,526 kcal MEn reported by Meyer et al. (1980a). The protein need for egg production has been observed to vary from 10 to
18 percent of the diet, with the value of 14 percent chosen as being the most representative.
Research on the amino acid requirements of turkeys has largely been conducted on the starting poult. With the exception
of lysine and the sulfur amino acids, little experimentation has been done to determine the amino acid requirements of
growing turkeys. Fisher (1982a) and Hurwitz et al. (1983a) employed body analyses and feed intake together with calculated
maintenance needs to estimate requirements. The protein requirements shown in Table 3-1 are based on either actual
experimentation, modeling, or are calculated as a ratio with lysine when the requirement for lysine at the ages in question has
been measured experimentally.
The starting poult's arginine requirement of 1.6 percent of the diet is supported by the research of Almquist (1952) and
Warnick and Anderson (1973) and the modeling of Hurwitz et al. (1983a). Dunkelgod et al. (1970) and D'Mello and Emmans
(1975) reported higher arginine requirement
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NUTRIENT REQUIREMENTS OF TURKEYS
38
values when they fed amino acid mixtures or diets based on wheat-corn gluten meal, respectively.
The isoleucine requirement listed for starting turkeys (1 percent of the diet) is based largely on the research of Warnick
and Anderson (1973) and agrees well with the value of 1.03 percent obtained from modeling by Hurwitz et al. (1983a).
Similarly, the leucine requirement (1.9 percent of the diet) is based on the determined value of 1.86 percent reported by
Warnick and Anderson (1973) and 1.96 percent from modeling by Hurwitz et al. (1983a).
The lysine and sulfur amino acid needs have been well investigated because of their frequent limitation under practical
conditions. Starting poults require 1.6 percent lysine in the diet. This value represents an average of the determined values
1.55 percent (Balloun and Phillips, 1957b), 1.6 percent (Kummero et al., 1971), 1.68 percent (Warnick and Anderson, 1973),
1.5 percent (Tuttle and Balloun, 1974), and 1.55 percent (D'Mello and Emmans, 1975). The value of 1.42 percent obtained by
modeling (Hurwitz et al., 1973) is noticeably lower than those measured by bioassay. Lysine needs after the first 4 weeks of
life have been derived mainly from the research of Tuttle and Balloun (1974), Jensen et al. (1976), and Potter et al. (1981).
The poult's requirement of 0.55 percent methionine in the diet is greater than the 0.53 percent given in the previous
edition of this report and is the value that best represents the reports of Almquist (1952), Baldini et al. (1957), and Murillo
and Jensen (1976a). Requirement values beyond starting were provided from the experimentation of Murillo and Jensen
(1976a) and Behrends and Waibel (1980). The total sulfur amino acid requirement value of 1.1 percent for starting poults was
derived from the observations of 1.04 percent by Warnick and Anderson (1973), 1.05 percent by Murillo and Jensen (1976b),
1.10 percent by Potter and Shelton (1979), and 1.1 percent by Behrends and Waibel (1980), as well as the 1.05 percent from
modeling by Hurwitz et al. (1983a). Requirement values specifically for methionine subsequent to starting largely represent
the observed needs to optimize performance as reported by Potter and Shelton (1979, 1980), Murillo and Jensen (1976a), and
Behrends and Waibel (1980), together with the modeling estimate by Hurwitz et al. (1983a).
Minerals
The calcium requirement determined with starting poults has been reported to be as high as 1.7 percent (Motzok and
Slinger, 1948) and 1.5 percent (Wilcox et al., 1953) and as low as 1.0 percent (Slinger et al., 1961) and 0.81 percent (Formica
et al., 1962). Neagle et al. (1968) reported a requirement of 1.2 percent dietary calcium when total phosphorus and vitamin D
levels were 0.8 percent and 1,100 ICU/kg of diet, respectively. The latter calcium requirement for growing turkeys has been
substantiated by Nelson et al. (1961), Sullivan (1961), and Formica et al. (1962). Hens in egg production need approximately
2.25 percent calcium in the feed, as shown by Balloun and Miller (1964a), Arends et al. (1967), Potter et al. (1974), and
Waldroup et al. (1974b).
The nonphytate phosphorus requirement of 0.6 percent for starting poults agrees with the research reported by Almquist
(1954), Bailey et al. (1986), and Stevens et al. (1986). This value has been shown to decrease with age (Day and Dilworth,
1962; Sullivan, 1962). Reported nonphytate phosphorus requirements for breeder hens in egg production range from 0.3
percent (Waldroup et al., 1974b; Slaugh et al., 1989) to 0.55 percent (Atkinson et al., 1976). The latter relatively high value
probably occurred because of a low phosphorus availability in the feedstuffs employed; thus 0.35 percent was selected to
represent the requirement.
The magnesium requirement, given as 500 mg/kg of diet, has been reduced from the 600 mg listed in the previous
edition to better reflect the value of 475 mg/kg reported by Sullivan (1964). The manganese requirement may vary with the
type of diet and supplement used. The recommended value of 60 mg/kg is the same as the requirement observed by Kealy
and Sullivan (1966). The same level was reported by Atkinson et al. (1967b) as the requirement for breeder hens. Zinc needs
are known to depend on the levels of other dietary constituents. The recommended level of 70 mg/kg was determined with
practical diets having phytic acid present, whereas 41 mg/kg were adequate in a purified diet where phytic acid was absent
(Dewar and Downie, 1984).
Vitamins
The previous requirement for vitamin A was listed as 4,000 IU/kg of diet. Vitamin A at 5,000 IU/kg of feed provides for
maximum growth performance and liver storage (Prinz et al., 1986) and has been chosen to represent the requirement,
although 2,000 IU/kg will also support optimal performance (Prinz et al., 1983). Vitamin A at 5,000 IU/kg is also
recommended for breeder hens, but lower levels (about 2,500 IU/kg) have been shown to maintain egg production,
hatchability, and survival (Stoewsand and Scott, 1961; Jensen et al., 1965).
Vitamin D3 at 900 IU/kg of feed has been shown to be more than adequate for the starting poult in most studies (Baird
and Greene, 1935; Hammond, 1941; Stadelman et al., 1950); however, Neagle et al. (1968) found that 1,100 IU/kg was
necessary to maximize both growth and toe ash concentration when the diet contained 1.2 percent calcium and 0.8 percent
total phosphorus. Discrepancies in vitamin D3 needs of poults
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NUTRIENT REQUIREMENTS OF TURKEYS
39
may relate to the level of this vitamin in the breeder hen's feed. Stevens et al. (1984) observed that 900 IU/kg in the breeder
hen's diet supported maximum egg yield, hatchability, and subsequent survival of the poult, but liver storage was considered
marginal.
The value given as the vitamin E requirement of starting turkeys is the same as that reported by Scott et al. (1965) when
the dietary selenium concentration was 0.1 mg/kg. The vitamin E requirement of breeder hens was observed to be twice this
level (24 IU/kg; Jensen and McGinnis, 1957). Extensive increases in vitamin E well above requirements for optimal growth
are necessary in order to provide the carcass meaningful protection against oxidative rancidity when carcasses are held in
frozen storage (Sheldon, 1984).
All other vitamin requirements have been determined only for the first 4 or 8 weeks of age. In some instances, there is
good agreement among the researchers on the requirement value but, in other instances, considerable disparity exists. The
committee has revised the requirement values given for several vitamins either to better represent old information or to reflect
new reports. Vitamin K at 1 mg/kg of diet was increased to 1.75 mg/kg to be the same as the value observed by Griminger
(1957) to optimize blood prothrombin time. The new value is considered adequate under practical conditions because poults
used by Griminger (1957) were reared in wire-floored pens and coprophagy, as an additional source of vitamin K, was
prevented.
Ruiz and Harms (1989a) reported that the poult's requirement for riboflavin was greater than 3.5 mg/kg of diet. The
value given in the previous edition was 3.6 mg/kg, and this has been increased to 4.0 mg/kg. Conversely, Ruiz and Harms
(1989b) reported the pantothenic acid requirement to be less than 8.6 mg/kg of diet; thus the previously listed requirement of
11 mg/kg was reduced to 10 mg/kg.
The dietary need for choline is known to be influenced by the levels of other nutrients involved in methyl group
metabolism. The previously listed choline requirement was 1,900 mg/kg of diet, which was largely based on the report of
Evans (1943), wherein the levels of ancillary nutrients influential to methyl group metabolism were not ensured. Harms and
Miles (1984) reported that the choline requirement for poults between 0 and 4 weeks of age was less than 1,490 mg/kg of
diet. Blair et al. (1986), using turkeys between 4 and 8 weeks of age, reported that the requirement was less than 1,250 mg/kg.
To reflect these observations, the present requirement has been reduced to 1,600 and 1,400 mg/kg of diet for the period from
0 to 4 and 4 to 8 weeks, respectively.
The requirements for many vitamins after 8 weeks of age have not been determined for turkeys. Only measurements of
the vitamin D3, pantothenic acid, biotin, and folacin requirements have been conducted on breeder hens.
TABLE 3-3 Body Weights and Feed Consumption of Large-Type Turkeys during the Holding and Breeding Periods
Females
Males
Age (weeks)
Weight (kg)
Egg Production (%)
Feed per Turkey Daily (g)
Weight (kg)
Feed per Turkey Daily (g)
20
8.4
0
260
14.3
500
25
9.8
0
320
16.4
570
a
310
19.1
630
30
11.1
0
35
11.1
68
280
20.7
620
40
10.8
64
280
21.8
570
45
10.5
58
280
22.5
550
50
10.5
52
290
23.2
560
55
10.5
45
290
23.9
570
10.6
38
290
24.5
580
60
NOTE: These values are based on experimental data involving ''in-season" egg production (that is, November through July) of
commercial stock. It is estimated that summer breeders would produce 70 to 90 percent as many eggs and consume 60 to 80 percent as
much feed as in-season breeders.
a Light stimulation is begun at this point.
Requirement values for other vitamins were estimated from experimentally determined values for younger ages and
changes in requirements observed with chickens.
TURKEY BREEDERS
Through the first 12 to 16 weeks of age, male and female turkeys being grown for reproductive purposes generally have
been fed the same diet as birds intended for meat production. Thereafter, various efforts have been implemented to avoid
obesity. Limiting body weight gain of males by either restricting feed access (Krueger et al., 1978) or providing a low-protein
feed for ad libitum consumption (Meyer et al., 1980b) is effective as long as the practices are not so severe that they delay
semen production. Typical nutrient levels employed from this time through the active breeder period correspond to those of
the holding feed, as given in Table 3-1.
Excess body weight of hens is less of a problem than with males because an extensive loss of body weight occurs with
hens as time in lay progresses. Table 3-3 includes a sample of hen performance through the breeder period. Inadequate body
weight gain prior to stimulatory lighting delays the onset of lay and reduces egg production (Krueger et al., 1978; Meyer et
al., 1980a). Starting both sexes on feed having the lowest concentration of nutrients for which a balance can be formulated
and continuing this regimen to and through the breeder period on an ad libitum consumption basis minimizes the likelihood
of obesity without adversely affecting performance (Ferket and Moran, 1985, 1986).
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NUTRIENT REQUIREMENTS OF GEESE
40
4
Nutrient Requirements of Geese
Geese are reared under a variety of feeding programs. In the production of "farm geese," the goslings are given starter
feed for about 2 weeks and then allowed to forage for a variety of pasture and grain feedstuffs. Under these conditions, they
are marketable at about 18 weeks. In another program, the goslings are fed limited amounts of prepared feed throughout the
growing period but are still allowed considerable foraging. These geese are marketed at about 14 weeks of age, following
liberal feeding of a high-energy finishing diet. Geese may also be provided feed for ad libitum consumption in confinement
and marketed as "junior" or "green geese" at about 10 weeks. A program practiced in European countries involves the
production of goose livers for paté de foie gras. The geese are grown to about 12 weeks and are then force-fed a high-energy
diet for the production of livers of high-fat content. Geese for breeding purposes are fed holding and breeding diets for the
intensive production of fertile eggs.
The nutrient requirements data presented in Table 4-1 are primarily applicable to geese reared in confinement. The
nitrogen-corrected metabolizable energy (MEn) concentrations heading each column are not requirements; instead they
represent what are considered typical dietary MEn values used for rearing geese commercially. Feed consumption by growing
geese decreases as dietary MEn level increases, but not in direct proportion (Stevenson, 1985). Consequently, geese fed highenergy diets consume greater amounts of energy, and deposit more body fat, than do geese fed lower-energy diets (Roberson
and Francis, 1963a; Stevenson, 1985).
Data obtained from research done since 1980 by using fast-growing geese were used to establish the protein
requirements given in Table 4-1. These data show that starting geese (0 to 4 weeks of age) require no more than 20 percent
protein (Allen, 1981; Nitsan et al., 1983; Summers et al., 1987) for satisfactory growth, carcass composition, and feathering.
Earlier research (Roberson and Francis, 1963a,b) with White Chinese geese had indicated that the protein requirement during
the period from 0 to 6 weeks was 24 percent. In view of recent data, it is questionable whether this higher requirement applies
to modern, commercial geese. No research data on the protein requirement of geese used for breeding or egg production were
found in the literature.
Little information has been published describing the amino acid, mineral, or vitamin requirements of geese (Appendix
Table A-5). Roberson and Francis (1966) reported that 0.90 percent lysine was needed for maximum growth and efficiency of
feed utilization by 0- to 3-week-old White Chinese geese fed a diet containing
TABLE 4-1 Nutrient Requirements of Geese as Percentages or Units per Kilogram of Diet (90 percent dry matter)
Unit
0 to 4 Weeks; 2,900a
After 4 Weeks; 3,000a
Breeding; 2,900a
Nutrients
Protein and amino acids
Protein
%
20
15
15
Lysine
%
1.0
0.85
0.6
Methionine + cystine
%
0.60
0.50
0.50
Macrominerals
Calcium
%
0.65
0.60
2.25
Nonphytate phosphorus
%
0.30
0.3
0.3
Fat soluble vitamins
A
IU
1,500
1,500
4,000
IU
200
200
200
D3
Water soluble vitamins
Choline
mg
1,500
1,000
?
Niacin
mg
65.0
35.0
20.0
Pantothenic acid
mg
15.0
10.0
10.0
mg
3.8
2.5
4.0
Riboflavin
NOTE: For nutrients not listed or those for which no values are given, see requirements of chickens (Table 2-5) as a guide. Where
experimental data are lacking, values typeset in bold italic represent an estimate based on values obtained for other ages or species.
a These are typical dietary energy concentrations expressed in kcal ME /kg diet.
n
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NUTRIENT REQUIREMENTS OF GEESE
41
20 percent protein and 2,950 kcal MEn/kg. More recently, Mateova et al. (1980) found that 1.10 percent lysine was
satisfactory for starting geese. Mateova et al. (1980) also reported that from 4 to 8 weeks of age geese needed 0.85 percent
lysine in a diet containing 2,945 kcal MEn/kg. Nitsan et al. (1983) used body composition, maintenance needs, and absorption
rate of amino acids to estimate the lysine requirements of geese. Subsequent testing of the results in feeding trials indicated
that goslings required 1.07 and 0.60 percent lysine during the period from 0 to 2 and 2 to 7 weeks, respectively. Requirements
of geese for other essential amino acids were estimated by Nitsan et al. (1983), and the results indicated that 0.58 percent total
sulfur amino acids (TSAA) and 0.29 percent methionine were needed from 0 to 2 weeks of age and 0.47 percent TSAA and
0.15 percent methionine were required from 2 to 7 weeks.
Calcium and total phosphorus requirements of geese were estimated at 0.4 percent and 0.46 percent of the diet,
respectively, for geese from 0 to 4 weeks of age (Aitken et al., 1958). These estimates have not been corroborated by recent
research. Briggs et al. (1953) documented the need for dietary folic acid, choline, and niacin by goslings but did not estimate
requirements. Battig et al. (1953) reported that 66 mg of dietary niacin per kilogram of diet (40 mg supplemented plus 26 mg
in the ingredients) were required to prevent perosis and maximize growth of geese to 3 weeks of age.
Serafin (1981) fed purified diets to Embden goslings from hatch to 2 or 3 weeks and found that, for growth and
liveability, requirements for riboflavin, niacin, pantothenic acid, and choline were no more than 3.8, 31.2, 12.6, and 1,530 mg/
kg, respectively. Laboratory analysis of the basal purified diet showed that concentrations of the vitamins studied were very
low; hence the requirement data reported herein represent levels of supplemental vitamins that were supplied in highly
available forms. Thus, supplemental vitamins, which probably were readily utilized by the geese, were used to establish the
requirements for riboflavin, niacin, pantothenic acid, and choline. Requirements established in this way may not be totally
applicable to feeding commercial geese because vitamins supplied by commonly used ingredients of geese diets are less
available than those of supplemental origin.
TABLE 4-2 Approximate Body Weights and Feed Consumption of Commercially Reared Male and Female Geese to 10 Weeks of Age
Average Body Weight (kg)
Feed Consumption by 2-Week Period (kg)
Cumulative Feed Consumption (kg)
Age (weeks)
0
0.11
0.00
0.00
2
0.82
0.96
0.96
4
2.05
2.93
3.89
6
3.05
3.20
7.09
8
4.05
4.34
11.43
4.85
4.68
16.11
10
The paucity of research on the nutrient requirements of geese illustrates the need for additional efforts focused on this
area of nutrition.
Body weight and feed consumption data presented in Table 4-2 are approximations obtained from a combination of
research results and input from persons involved in the production of geese.
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NUTRIENT REQUIREMENTS OF DUCKS
42
5
Nutrient Requirements of Ducks
Ducks can be grown successfully in either of two environments—an open rearing system, in which the growing house
opens to an exercise yard with water for wading or swimming, or a confinement growing system, in which ducks are raised in
environmentally controlled houses with litter or combination litter and wire floors.
Pelleted diets are utilized more efficiently by ducks than are diets in mash form primarily because of reduced wastage
and ease of consumption (Wilson, 1973; Dean, 1986). Starter diets (0 to 2 weeks) usually are fed as pellets of 3.18 mm (1/8
inch) diameter, and grower diets (after 2 weeks) are given in 4.76-mm (3/16 inch) form (Elkin, 1987).
Ducks typically are given 2 or 3 feeds during the growing period. Information presented in Table 5-1 is on the basis of a
two-feed program; a diet containing 22 percent protein for the period of 0 to 2 weeks and a 16 percent protein diet for the
period from 2 to 7 weeks (Dean, 1972a, 1986). The need for 22 percent protein during the starting period, however, is
questionable because Wilson (1975) and Siregar et al. (1982) reported that protein levels of 18 and 19 percent, respectively,
in diets providing 3,000 to 3,025 kcal MEn/kg, were adequate from 0 to 2 weeks. A typical three-feed program may consist of
diets containing 20, 18, and 16 percent protein for the periods from 0 to 2, 2 to 4, and 4 to 7 weeks, respectively. The growth
rate of ducklings is not affected greatly by the MEn concentration of the diet; however, feed efficiency is usually improved
and carcass fat increased when dietary MEn is increased (Wilson, 1975; Leclercq, 1986). Few data are available documenting
the MEn values of feed ingredients for ducks. Mohamed et al. (1984) found that the MEn values of several feedstuffs were
very similar for ducks and broiler chickens.
Although most ducks grown commercially in the United States are White Pekins, considerable research
TABLE 5-1 Nutrient Requirements of White Pekin Ducks as Percentages or Units per Kilogram of Diet (90 percent dry matter)
Unit
0 to 2 Weeks; 2,900a
2 to 7 Weeks; 3,000a
Breeding; 2,900a
Nutrient
Protein and amino acids
Protein
%
22
16
15
Arginine
%
1.1
1.0
Isoleucine
%
0.63
0.46
0.38
Leucine
%
1.26
0.91
0.76
Lysine
%
0.90
0.65
0.60
Methionine
%
0.40
0.30
0.27
Methionine + cystine
%
0.70
0.55
0.50
Tryptophan
%
0.23
0.17
0.14
Valine
%
0.78
0.56
0.47
Macrominerals
Calcium
%
0.65
0.60
2.75
Chloride
%
0.12
0.12
0.12
Magnesium
mg
500
500
500
Nonphytate phosphorus
%
0.40
0.30
Sodium
%
0.15
0.15
0.15
Trace minerals
?
Manganese
mg
50
?b
Selenium
mg
0.20
?
?
Zinc
mg
60
?
?
Fat soluble vitamins
A
IU
2,500
2,500
4,000
IU
400
400
900
D3
E
IU
10
10
10
K
mg
0.5
0.5
0.5
Water soluble vitamins
Niacin
mg
55
55
55
Pantothenic acid
mg
11.0
11.0
11.0
Pyridoxine
mg
2.5
2.5
3.0
mg
4.0
4.0
4.0
Riboflavin
NOTE: For nutrients not listed or those for which no values are given, see requirements of broiler chickens (Table 2-5) as a guide. Where
experimental data are lacking, values typeset in bold italics represent an estimate based on values obtained for other ages or species.
a These are typical dietary energy concentrations as expressed in kcal ME /kg diet.
n
b Question marks indicate that no estimates are available.
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NUTRIENT REQUIREMENTS OF DUCKS
43
data obtained by using other breeds of ducks (that is, Muscovy and "mule" ducks) have been used to fill several voids in the
requirement data of Table 5-1, especially with respect to amino acids and minerals. Published research reviewed in Appendix
Table A-6 on lysine and total sulfur amino acid (TSAA) requirements indicates that values listed in the previous edition of
this report were too high (Jeroch and Hennig, 1965; Dean, 1967; Gazo et al., 1970; Leclercq and de Carville, 1977a,b; Adams
et al., 1983; Elkin et al. 1986). Adjustments were made accordingly. In addition, a tentative methionine requirement for
starting ducks (0.40 percent) is given on the basis of data reported by Elkin et al. (1986). Noteworthy is information
published recently by Elkin et al. (1988) showing that the relative value of the D-methionine isomer was 78 percent of that of
the L-isomer. Consequently, in instances where supplemental methionine is needed in duck diets, adjustments may be needed
in supplemental levels of the DL-methionine sources used.
Only single papers have been published documenting the requirements of starting ducks for arginine, tryptophan,
leucine, isoleucine, and valine (Chen and Shen, 1979; Wu et al., 1984; Yu and Shen, 1984). The values for these nutrients
listed in Table 5-1 must therefore be viewed as tentative. The same is true of the requirement values for breeding ducks
because relevant information is scarce (Cvetanov et al., 1969).
Research to determine the mineral and vitamin requirements of ducks has focused primarily on the starting period (0 to 2
or 3 weeks of age). In most instances, data on these nutrients are meager, and, with the exception of some research on dietary
selenium and niacin requirements, only one report has appeared in the literature since 1980. Leclercq et al. (1990) reported
that the calcium requirements of Muscovy ducks were 0.46 and 0.42 percent for age periods of 3 to 8 and 8 to 12 weeks,
respectively. No information has been published recently on the calcium requirements for modern-day Pekin ducks.
TABLE 5-2 Approximate Body Weights and Feed Consumption of White Pekin Ducks to 8 Weeks of Age
Age (weeks)
Body Weight (kg)
Weekly Feed Consumption (kg)
Cumulative Feed Consumption (kg)
Male
Female
Male
Female
Male
Female
0
0.06
0.06
0.00
0.00
0.00
0.00
1
0.27
0.27
0.22
0.22
0.22
0.22
2
0.78
0.74
0.77
0.73
0.99
0.95
3
1.38
1.28
1.12
1.11
2.11
2.05
4
1.96
1.82
1.28
1.28
3.40
3.33
5
2.49
2.30
1.48
1.43
4.87
4.76
6
2.96
2.73
1.63
1.59
6.50
6.35
7
3.34
3.06
1.68
1.63
8.18
7.98
3.61
3.29
1.68
1.63
9.86
9.61
8
Body weight and feed consumption data for ducks from time of hatching to 8 weeks of age are given in Table 5-2.
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NUTRIENT REQUIREMENTS OF RING-NECKED PHEASANTS, JAPANESE QUAIL, AND BOBWHITE QUAIL
44
6
Nutrient Requirements of Ring-Necked Pheasants, Japanese
Quail, and Bobwhite Quail
As was true for geese and ducks, little information is available on the nutrient requirements of the game birds that are
most frequently considered part of the poultry industry—Ring-necked pheasants, Japanese quail, and Bobwhite quail.
Although these species do not constitute a major share of the poultry industry, there are an increasing number of specialized
farms involved in their production.
RING-NECKED PHEASANTS
Information available on the nutrient requirements of the Ring-necked pheasant indicates that diets of relatively high
nutrient concentrations are needed during the starting period (Table 6-1). Protein and amino acid needs, where documented
(Appendix Table A-7), resemble those of turkeys. Also, pheasants are especially prone to leg disorders and abnormal feather
growth when certain key nutrients such as niacin, riboflavin, choline, manganese, and zinc are inadequate (Sunde and Bird,
1957; Scott et al., 1959). Pheasant chicks are especially vulnerable to undefined dietary factors that impair leg development,
and including extra zinc in diets has been shown to reduce the impact of these factors (Cook et al., 1984). A high level of
calcium, as in a breeder ration, can cause leg problems and high mortality if fed to pheasant chicks (Woodard et al., 1979).
All nutrient requirements listed for female pheasants in egg production except for protein are tentative. Data presented
by Monetti et al. (1982, 1985) indicate that dietary protein concentration should be maintained so that percentage of protein
per megacalorie MEn/kg of diet does not exceed 5.6.
Often, pheasants are fed diets designed to produce birds for use on game-release farms. Diets relatively high in protein
and low in energy may be used to encourage the development of lean pheasants suitable for release.
JAPANESE QUAIL
Japanese quail are used for commercial specialty meat and egg production and also are valued research animals.
Consequently, the nutrient requirements of Japanese quail have been documented to a greater extent than have those of other
game bird species. Few definitive data have been published since 1984, when the previous edition of this report was
published and
TABLE 6-1 Nutrient Requirements of Ring-Necked Pheasants as Percentages or Units per Kilogram of Diet (90 percent dry matter)
Unit
0 to 4 Weeks; 2,800a
4 to 8 Weeks; 2,800a
9 to 17 Weeks; 2,700a
Breeding; 2,800a
Nutrient
Protein and amino acids
Protein
%
28
24
18
15
Glycine + serine
%
1.8
1.55
1.0
0.50
Linoleic Acid
%
1.0
1.0
1.0
1.0
Lysine
%
1.5
1.40
0.8
0.68
Methionine
%
0.50
0.47
0.30
0.30
Methionine + cystine
%
1.0
0.93
0.6
0.60
Protein
%
28
24
18
15
Macrominerals
Calcium
%
1.0
0.85
0.53
2.5
Chlorine
%
0.11
0.11
0.11
0.11
Nonphytate phosphorus
%
0.55
0.50
0.45
0.40
Sodium
%
0.15
0.15
0.15
0.15
Trace minerals
Manganese
mg
70
70
60
60
Zinc
mg
60
60
60
60
Water soluble vitamins
Choline
mg
1,430
1,300
1,000
1,000
Niacin
mg
70.0
70
40.0
30.0
Pantothenic acid
mg
10.0
10.0
10.0
16.0
mg
3.4
3.4
3.0
4.0
Riboflavin
NOTE: Where experimental data are lacking, values typeset in bold italics represent an estimate based on values obtained for other ages
or species. For nutrients not listed or those for which no values are given, see requirements of turkeys (Table 3-1) as a guide.
a These are typical dietary energy concentrations, expressed in kcal ME /kg diet.
n
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NUTRIENT REQUIREMENTS OF RING-NECKED PHEASANTS, JAPANESE QUAIL, AND BOBWHITE QUAIL
Shim and Vohra (1984) presented a comprehensive
review. Data appearing since 1984 have supported the values
listed in the 1984 edition for protein (Sinha and Verma,
1984; Steigner, 1990) and for total sulfur amino acids
(TSAA; Shrivastav and Panda, 1987) for the starting and
growing period. In the instance of protein, however, Steigner
(1990) reported that a strain of Japanese quail selected for
rapid growth required a greater dietary protein concentration
than did random-bred quail. Similarly, information provided
by Shim and Lee (1984, 1988) and by Shim and Chen (1989)
showed that the dietary requirements for lysine and TSAA
for breeding quail in the 1984 edition were appropriate in
relation to the stated metabolizable energy contents of the
diet. The lack of data to further define requirements or to
corroborate single sets of observations (Appendix Table A-8)
on requirements of Japanese quail, especially breeding quail,
necessitates the continued listing of a large number of
tentative requirement values in Table 6-2.
TABLE 6-2 Nutrient Requirements of Japanese Quail (Coturnix)
as Percentages or Units Per Kilogram of Diet (90 percent dry
matter)
Nutrient
Unit
Starting and
Breeding;
Growing;
2,900a
2,900a
Protein and amino acids
Protein
%
24.0
20.0
Arginine
%
1.25
1.26
Glycine + serine
%
1.15
1.17
Histidine
%
0.36
0.42
Isoleucine
%
0.98
0.90
Leucine
%
1.69
1.42
Lysine
%
1.30
1.00
Methionine
%
0.50
0.45
Methionine +
%
0.75
0.70
cystine
Phenylalanine
%
0.96
0.78
Phenylalanine +
%
1.80
1.40
tyrosine
Threonine
%
1.02
0.74
Tryptophan
%
0.22
0.19
Valine
%
0.95
0.92
Fat
Linoleic acid
%
1.0
1.0
Macrominerals
Calcium
%
0.8
2.5
Chlorine
%
0.14
0.14
Magnesium
mg
300
500
Nonphytate
%
0.30
0.35
phosphorus
Potassium
%
0.4
0.4
Sodium
%
0.15
0.15
Trace minerals
Copper
mg
5
5
Iodine
mg
0.3
0.3
Iron
mg
120
60
Manganese
mg
60
60
Selenium
mg
0.2
0.2
Zinc
mg
25
50
Fat soluble vitamins
A
IU
1,650
3,300
ICU
750
900
D3
E
IU
12
25
K
mg
1
1
Water soluble vitamins
mg
0.003
0.003
B12
Biotin
mg
0.3
0.15
Choline
mg
2,000
1,500
Folacin
mg
1
1
Niacin
mg
40
20
Pantothenic acid
mg
10
15
Pyridoxine
mg
3
3
Riboflavin
mg
4
4
mg
2
2
Thiamin
45
growing periods, see requirements for turkeys (Table 3-1) as a
guide.
a These are typical dietary energy concentrations, expressed in
kcal MEn/kg diet.
TABLE 6-3 Nutrient Requirements of Bobwhite Quail as
Percentages or Units per Kilogram of Diet (90 percent dry matter)
Unit
0 to 6
After 6
Breeding;
Nutrient
Weeks;
Weeks;
2,800a
a
a
2,800
2,800
Protein and amino acids
Protein
%
26
20.0
24.0
Methionine
%
1.0
0.75
0.90
+ cystine
Fat
Linoleic acid
%
1.0
1.0
1.0
Macrominerals
Calcium
%
0.65
0.65
2.4
Nonphytate
%
0.45
0.30
0.70
phosphorus
Sodium
%
0.15
0.15
0.15
Trace minerals
Chlorine
%
0.11
0.11
0.11
Iodine
mg
0.30
0.30
0.30
Water soluble vitamins
Choline
mg
1,500.0
1,500.0
1,000.0
Niacin
mg
30.0
30.0
20.0
Pantothenic
mg
12.0
9.0
15.0
acid
mg
3.8
3.0
4.0
Riboflavin
NOTE: Where experimental data are lacking, values typeset in
bold italics represent an estimate based on values obtained for
other ages or species. For values not listed for the startinggrowing periods, see requirements for turkeys as a guide.
a These are typical dietary energy concentrations, expressed in
kcal MEn/kg diet.
Bobwhite Quail
The committee has made few changes in the nutrient
specifications for Bobwhite quail (Table 6-3). Its
reevaluation of the data (Appendix Table A-9) used to
establish the previous requirements resulted in some
modifications in protein, TSAA, calcium, and phosphorus
recommendations for starting-growing Bobwhite quail. As
with other game birds reared commercially, Bobwhite quail
grown for game-release farms should be fed diets of
relatively low energy content during the growing period to
prevent excessive fattening.
NOTE: Where experimental data are lacking, values typeset in
bold italics represent an estimate based on values obtained for
other ages or species. For values not listed for the starting-
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SIGNS OF NUTRITIONAL DEFICIENCIES IN CHICKENS AND TURKEYS
46
7
Signs of Nutritional Deficiencies in Chickens and Turkeys
Clinical manifestation of nutrient deficiencies often occurs in conjunction with an alteration of normal biological
processes that are unique for the nutrient. Some enzymes depend on particular vitamins and minerals for their functioning,
and their activity diminishes with an inadequacy. In other instances, a particular physiological response or change in
metabolite concentration may occur. This information was primarily obtained from formal experiments in which the
inadequacies were definitive. Under field conditions, nutrient inadequacies are usually marginal, occasionally multiple, and
often confounded with management problems or disease. To supplement physical observation of these signs, the committee
has provided biochemical and physiological measurements for use in diagnosis. Table 7-1 presents a summary of the known
biochemical and physiological measurements for diagnosing each nutrient deficiency. Additional information is available in
the associated references.
Inadequate dietary vitamins and minerals in the chicken or turkey hen's diet are likely to reduce the egg contents
accordingly and have adverse effects on embryonic development. Normal embryonic development proceeds through several
events at which death of the embryo is common. The largest number of deaths occur during the transition from anaerobic to
aerobic respiration with the establishment of the chorioallantois, which takes place between 3 to 4 days incubation and
emergence at 18 to 21 days incubation. The same problems occur with other poultry species, and nutrient inadequacies
generally accentuate death rates at these times (Couch and Ferguson, 1972).
Embryos are well developed at the end of incubation, and embryos that die as a result of nutrient deficiencies at this time
may exhibit typical physical symptoms. These symptoms are assembled for each nutrient in Table 7-2. The symptoms can be
similar for different nutrients, and the extent of the inadequacy may change the nature of the symptoms as well as when death
occurs. Deficiency symptoms are expressed to a greater extent in growing birds than in adults. Table 7-3 gives a list of these
symptoms by tissue affected, as a diagnostic aid. The table also presents information on these changes such that each can be
rationalized in terms of nutrient function. References provided are not complete but are intended to be salient and most recent
for cross-indexing purposes. Again, such information is usually the product of formal experimentation and not complicated
by practical circumstances.
PROTEIN AND AMINO ACID DEFICIENCIES
Protein is made up of amino acids. The need for the essential amino acids determines the need for protein, and a
reduction in dietary protein that results in deficiencies of several essential amino acids creates general symptoms. Productive
activities suffer the most. For example, the energy used by growing birds is heavily committed to assembling the contractile
elements in muscle cells but not to increasing cell number; thus protein inadequacies readily affect muscle size but not fiber
number (Timson et al., 1983). Similarly, the effect of protein inadequacies on protein synthesis in the liver and oviduct is
greatest with the laying hen (Muramatsu et al., 1987).
Deficiencies of individual essential amino acids usually have the same effect as when protein is deficient; however,
additional symptoms may appear that characterize certain amino acids. Inadequate lysine is known to cause depigmentation
of the wing feathers in Bronze turkey poults (Vohra and Kratzer, 1959) and certain colored chicks (Klain et al., 1957). A
variety of abnormalities in feather development occur with deficiencies of arginine, valine, leucine, isoleucine, tryptophan,
phenylalanine, and tyrosine in growing chicks (Newberne et
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SIGNS OF NUTRITIONAL DEFICIENCIES IN CHICKENS AND TURKEYS
TABLE 7-1 Biochemical and Physiological Measurements for Diagnosis of Nutrient Deficiencies in Chickens and Turkeys
Nutrients
Biochemical and Physiological Measurements
References
Histidine
Reduced breast muscle anserine and carnosine.
Robbins et al., 1977; Amend et al., 1979
Lysine
Reduced hemoglobin and hematocrit.
Braham et al., 1961
Rogers, 1969; Nockels and Phillips, 1971;
Vitamin A
Hepatic vitamin A is indicative of a deficiency, but blood
Jensen, 1974; Bruckental and Ascarelli, 1975;
level is not. Liver xanthine dehydrogenase and kidney
Nockels et al., 1984
arginase both increase even in the first stages of a deficiency.
Reduced glycogen phosphorylase in liver, and red and white
muscles. Increased thyroid size and reduced T3 and T4.
Vitamin D
Calcium-binding protein of intestine; 1,25-(OH)2-D3 versus
Bar et al., 1972; Ohmdahl and DeLuca, 1973;
24,25-(OH)2-D2 in serum (complicated by dietary calcium
Morrissey et al., 1977; Boyan and Ritter, 1984;
and phosphorus); plasma alkaline phosphatase;
Kaetzel and Soares, 1985
nonproteolipid phospholipid content of rachitic cartilage.
Vitamin E
Superoxide dismutase; glutamic-oxaloacetictransaminase;
Walter and Jensen, 1964; Arnold et al., 1974;
plasma and tissue vitamin E concentration (all measurements
Sklan et al., 1981; Sklan and Donoghue, 1982
affected by selenium as well).
Vitamin K
Prothrombin clotting time of plasma.
Griminger et al., 1970
Thiamin
Transketolase in erythrocytes and leucocytes; plasma pyruvic
Lofland et al., 1963; Anonymous, 1977
acid.
Riboflavin
Liver xanthine dehydrogenase; erythrocyte glutathione
Chou, 1971; Lee, 1982
reductase.
Niacin
Level and ratio of niacin excretion products N'-methylDarby et al., 1975
nicotinamide and N'-methyl-2-pyridone-5-carboxyamide
(untested for fowl).
Biotin
Blood pyruvate carboxylase; ratio of C 16:1 to C 18:0 fatty
Edwards, 1974; Whitehead and Bannister, 1980
acids in blood.
Pantothenic acid
Hepatic coenzyme A.
Cupo and Donaldson, 1986
Pyridoxine
Serum glutamic oxaloacetic transaminase; plasma glycineDaghir and Balloun, 1963; Sifri et al., 1972; Lee
et al., 1976
serine ratio aspartic aminotransferase.
Folacin
Dihydrofolic acid reductase in liver; serine hydroxymethyl
Rabbani et al., 1973; Zamierowski and Wagner,
transferase in liver.
1977
B12 in blood; excretion of methylmalonic acid.
Cox and White, 1962; Lau et al., 1965
Vitamin B12
Choline
Serum phospholipids.
Seifter et al., 1972
Linoleic acid
Linoleate, arachidonate, and eicosatrienoate concentrations in
Machlin and Gordon, 1960
liver lipids.
Bar et al., 1972, 1978a,b; Bar and Hurwitz, 1973
Calcium
Calcium in hen's blood (but not in chick's unless deficiency is
severe); intestinal calcium-binding protein (complicated by
D3 metabolites and phosphorus); turkey poults differ from
chicks.
Chlorine
Hemoconcentration; alkalosis.
Leach and Nesheim, 1963; Cohen and Hurwitz,
1974; Hamilton and Thompson, 1980
Copper
Plasma ceruloplasmin; lysyl oxidase in aorta, liver, tendon,
Kim and Hill, 1966; Miller and Stake, 1974;
and bone; erythrocyte superoxide dismutase.
Bettger et al., 1979; Opsahl et al., 1982
Iodine
Plasma thyroxine and tri-iodothyronine.
Singh et al., 1968
Iron
Hematocrit; blood hemoglobin concentration; transferrin
Davis et al., 1962; Waddell and Sell, 1964;
saturation; anemia with lipemia.
Planas, 1967
Magnesium
Magnesium concentration in blood.
Sell et al., 1967; Hajj and Sell, 1969
Manganese
Chondroitin sulfate in bone; manganese concentration in
Leach, 1968; Reid et al., 1973; DeRosa et al., 1980
bone; superoxide dismutase.
Phosphorus
Serum inorganic phosphorus; renal calcium-binding protein.
Miller and Stake, 1974; Bar et al., 1978a,b
Potassium
Plasma potassium; metabolic acidosis (complicated by
Burns et al., 1953; Cohen and Hurwitz, 1974
sodium).
Selenium
Plasma glutathionine peroxidase.
Noguchi et al., 1973; Dean and Combs, 1981;
Cantor et al., 1982
Sodium
Metabolic acidosis (complicated by potassium).
Nott and Combs, 1969; Cohen and Hurwitz, 1974
Plasma and bone zinc; thymidine kinase; alkaline
Miller and Stake, 1974; Oberleas and Prasad,
Zinc
phosphatase and collagenase in bone.
1974; Starcher et al., 1980; Bettger et al., 1979
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47
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SIGNS OF NUTRITIONAL DEFICIENCIES IN CHICKENS AND TURKEYS
TABLE 7-2 Signs of Deficiency in the Embryo
Nutrients
Deficiency Signs
Vitamin A
Death at about 48 hours of incubation from failure to
develop the circulatory system; abnormalities of kidneys,
eyes, and skeleton.
Vitamin D
Death at about 18 or 19 days of incubation, with
malpositions, soft bones, and with a defective upper
mandible prominent.
Vitamin E
Early death at about 84 to 96 hours of incubation, with
hemorrhaging and circulatory failure (implicated with
selenium).
Vitamin K
No physical deformities from a simple deficiency, nor can
they be provoked by antivitamins, but mortality occurs
between 18 days and hatching, with variable hemorrhaging.
Thiamin
High embryonic mortality during emergence but no
obvious symptoms other than polyneuritis in those that
survive.
Riboflavin
Mortality peaks at 60 hours, 14 days, and 20 days of
incubation, with peaks prominent early as deficiency
becomes severe. Altered limb and mandible development,
dwarfism, and clubbing of down are defects expressed by
embryo.
Niacin
Embryo readily synthesizes sufficient niacin from
tryptophan. Various bone and beak malformations occur
when certain antagonists are administered during
incubation.
Biotin
High death rate at 19 to 21 days of incubation, and
embryos have parrot beak, chondrodystrophy, several
skeletal deformities, and webbing between the toes.
Pantothenic acid
Deaths appear around 14 days of incubation, although
marginal levels may delay problems until emergence.
Variable subcutaneous hemorrhaging and edema; wirey
down in poults.
Pyridoxine
Early embryonic mortality based on antivitamin use.
Folic acid
Mortality at about 20 days of incubation. The dead
generally appear normal, but many have bent tibiotarsus,
syndactyly, and mandible malformations. In poults,
mortality at 26 to 28 days of incubation with abnormalities
of extremities and circulatory system.
Mortality at about 20 days of incubation, with atrophy of
Vitamin B12
legs, edema, hemorrhaging, fatty organs, and head
between thighs malposition.
Manganese
Peak deaths prior to emergence. Chondrodystrophy,
dwarfism, long bone shortening, head malformations,
edema, and abnormal feathering are prominent.
Zinc
Deaths prior to emergence, and the appearance of
rumplessness, depletion of vertebral column, eyes
underdeveloped, and missing limbs.
Copper
Deaths at early blood stage with no malformations.
Iodine
Prolongation of hatching time, reduced thyroid size, and
incomplete abdominal closure.
Iron
Low hematocrit; low blood hemoglobin; poor extraembryonic circulation in candled eggs.
High incidence of dead embryos early in incubation.
Selenium
48
References
Asmundson and Kratzer, 1952; Thompson et al.,
1965; Heine et al., 1985
Sunde et al., 1978; Narbaitz and Tsang, 1989
Card et al., 1930; Latshaw and Osman, 1974
Griminger, 1964; Hauschka and Reid, 1978a
Polin et al., 1962; Charles et al., 1972
Romanoff and Bauernfeind, 1942; Landauer, 1967
Snell and Quarles, 1941; Landauer, 1956; Caplan,
1972
Cravens et al., 1994; Couch et al., 1947
Kratzer et al., 1955; Beer et al., 1963
Landauer, 1967
Sunde et al., 1950a; Kratzer et al., 1956a
Olcese et al., 1950; Ferguson et al., 1955
Lyons and Insko, 1937
Kienholz et al., 1961; Turk, 1965
Bird et al., 1963
Rogler et al., 1959a, b
Dewar et al., 1974; Morck and Austic, 1981
Latshaw et al., 1977
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SIGNS OF NUTRITIONAL DEFICIENCIES IN CHICKENS AND TURKEYS
TABLE 7-3 Nutrients Associated with Various Signs of Deficiency in Growing Birds
Deficiency Signs
Descriptions
Species
Skin lesions
Crusting and scab formation
Chick, poult,
around eyes and beak
Chick, poult
Bottoms of feet rough and
calloused with hemorrhagic
cracks
Scaliness on feet
Chick
Lesions around eyes, eyelids
Chick, poult
stuck together
Mouth, inflammation of oral
Poult, chick
mucosa (chicken black tongue)
Feather abnormalities
Uneven feather growth,
Chick, poult
abnormally long primary
feathers, feathers not lying
smoothly
Frizzled and rough
Chick, poult
Nervous disorders
Convulsions with hyperexcitability
Hyperirritability
Characteristic fright reaction with
tetanic spasms
Spastic cervical paralysis, neck
extended with birds appearing to
look down
Curled-toe paralysis, gross
enlargement of sciatic and
brachial nerves with myelin
degeneration
Encephalomalacia, tetanic spasms
with head retraction, hemorrhagic
lesions in cerebellum
Blood and vascular system
Macrocytic
Macrocytic, hyperchromic
Microcytic, hypochromic
Microcytic
Hemorrhage, intramuscular,
subcutaneous, internal from aortic
rupture
Exudative diathesis
Enlarged heart
Muscle
Bone disorders
Associated Nutrients
Biotin, pantothenic acid
Biotin, pantothenic acid
Zinc, niacin
Vitamin A
Niacin
Protein, amino acid
imbalance
Black pigmentation in breeds
with red and brown feathers
Depigmentation
Convulsions with head
retraction
Chick, poult, duckling
Chick, poult, duckling
Chick
Chick
Zinc, niacin, pantothenic
acid, folic acid, lysine
Vitamin D
Chick, poult,
Chick, pigeon
Copper, iron, folacin
Thiamin
Poult
Folacin
Chick
Riboflavin
Chick
Vitamin E
Anemia
All poultry
Vitamin B12
Folacin
Iron, copper
Pyridoxine
Vitamin K, copper
Chick, poult
Pyridoxine
Magnesium, sodium chloride
Chloride
Chick, poult
Chick, poult
Muscular dystrophy, white
areas of degeneration in
skeletal muscle
Cardiac myopathy
Gizzard myopathy
Soft, easily bent bones and
beak (rickets)
Selenium, vitamin E
Copper
Chick, duck, poult
Hock enlargement
Poult, chick, gosling,
duckling
Chick, poult
Perosis
Bowed legs
Shortening and thickening of
leg bones
Curled toes
Diarrhea
49
Poult
Poult
All poultry
Vitamin E, selenium
Vitamin E, selenium
Vitamin E, selenium
Vitamin D, calcium or
phosphorus deficiency or
imbalance
Niacin, zinc
Duck
Chick
Biotin, choline, vitamin
B12, manganese, zinc,
folacin
Niacin
zinc, manganese
Chick
Chick, duck, poult
Riboflavin
Niacin, riboflavin, biotin
NOTE: Slow growth and general lack of vigor are generally associated with malnutrition. The signs listed in this table are more specific
indications of deficiencies of particular nutrients.
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SIGNS OF NUTRITIONAL DEFICIENCIES IN CHICKENS AND TURKEYS
50
al., 1960; Robel, 1977; Penz and Kratzer, 1984). Chavez and Kratzer (1974) observed a foot pad dermatitis in poults
when methionine was deficient, but cystine had to be adequate for the dermatitis to occur. Grau (1945) reported a tongue
deformity in chicks fed a purified diet deficient in leucine, isoleucine, or phenylalanine, but these observations were not
confirmed by Bragg (1953) with practical feedstuffs.
VITAMIN DEFICIENCIES
Vitamin A
Substitution of the body's secretory epithelia by keratinized surfaces is the most important change occurring with a
vitamin A deficiency. Corneal, conjunctival, esophageal, and tracheal secretory membranes are all altered in chickens
(Aydelotte, 1963). Mucus formation depends on vitamin A (DeLuca et al., 1971). Loss of membrane integrity, in turn, alters
water retention (Lopen et al., 1973) and impairs the ability to withstand infection (Singh and Donovan, 1973; Sijtsma et al.,
1989). Inadequate vitamin A also reduces the immune system's response to challenge and further contributes to disease
susceptibility (Davis and Sell, 1989; Sklan et al., 1989).
The appearance of keratinized secretory surfaces is followed by a typical ataxia. Alterations in bone growth create
several areas of compression on the central nervous system that cause a loss in mobility (Howell and Thompson, 1967).
Inadequate vitamin A also adversely affects the pituitary-gonadal axis to create other symptoms that are not readily obvious
(Fletcher, 1971). Nockels et al. (1984) reported that hypothyroidism is an early indication of vitamin A deficiency in chicks.
Reductions in testes size, circulating testosterone, and fertility have been reported during vitamin A deficiency in cockerels
(Padedes and Garcia, 1959; Hall et al., 1980).
Muscles in vitamin-A-deficient birds have a high level of glycogen, which cannot be readily used because
phosphorylase activity is inordinately low (Nockels and Phillips, 1971; Sundeen et al., 1980). Alternatively, glucose is
provided by extensive gluconeogenesis from protein (Nir and Ascarelli, 1967; Bruckental et al., 1974), and nitrogen end
products increase such that deposits of uric acid appear in the kidneys and ureters (Bruckental and Ascarelli, 1975; Chandra
et al., 1984).
Vitamin A in feedstuffs is labile, and concentrated supplements are normally given to ensure that the requirement is met.
Misuse of these concentrates has led to occasional toxicosis problems. Skin lesions at the commissure of the beak, nose, and
eyes attributable to mucus membrane hyperplastic activity have been shown to occur in chicks within 72 hours after oral
dosing with 60,000 IU (Kriz and Holman, 1969). The appearance of rachitic bones together with a hyperplastic parathyroid
results from the antagonism known to exist with vitamin D (Metz et al., 1985; Tang et al., 1985; Veltmann et al., 1987).
Excessive vitamin A has also been shown to antagonize vitamin E (Vahl and Van't Klooster, 1987) and increase the
likelihood of a deficiency when vitamin E and selenium nutriture is marginal (Combs, 1976).
Plant source feedstuffs usually provide carotenoid pigments that may be converted into vitamin A. The most favorable
such pigment in this respect is β-carotene (Flegal et al., 1971), and conversion largely occurs at the intestine during
absorption (Sklan, 1983). Because of the susceptibility of vitamin A sources to oxidative losses, synthetic antioxidants often
are included in premixes and complete feeds (Grundboeck et al., 1977).
Vitamin D
Poultry require vitamin D to effectively use calcium. After absorption, the vitamin is hydroxylated at the 25-position in
the liver and then transferred to the kidney, where the 1,25-dihydroxy metabolite is formed (Ameenuddin et al., 1985). All of
the vitamin metabolites affect calcium utilization in one way or another, but the 1,25-dihydroxy-vitamin D seems to have the
greatest impact. Vitamin D metabolites induce the synthesis of calcium-binding proteins in the intestine, kidney, and uterus
through the efforts of vitamin D metabolites at both transcriptional and post-transcriptional levels. Calcium-binding proteins
enhance calcium absorption from the intestine, recovery from the urine, and shell deposition, respectively (Coty, 1980; Jande
et al., 1981; Roth et al., 1981; Clemens et al., 1988).
Vitamin D also induces the formation of osteocalcin, a protein in bone (Anonymous, 1981). Osteocalcin is believed to
participate in the organic-inorganic matrix. Vitamin D is implicated by converting specific glutamic acid residues in
osteocalcin to γ-carboxylglutamic acid metabolites that interact with calcium. Bone alterations associated with osteocalcin
appear to be more involved with resorption and turnover when calcium is needed elsewhere in the body than growth.
Presumably, vitamin D also provides proliferative signals for undifferentiated cells in the intestine (Cross and Peterlik, 1983)
and pancreatic islets (Clark et al., 1987).
Vitamin D2 represents the plant source of this vitamin and arises from the ultraviolet irradiation of ergosterol (Kobayashi
and Yasumura, 1973), whereas vitamin D3 occurs in animals upon irradiation of 7-dehydro-cholesterol in skin (Beadle,
1977). Vitamin D3 is about 10-fold more effective with chicks than vitamin D2 (Hurwitz et al., 1967). A large part of this
difference in
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51
activity seems to involve metabolite formation in the liver, where enhanced glucuronidation of the 25-hydroxy-vitamin D2
favors biliary excretion (Le Van et al., 1981).
Gross symptoms occurring because of a vitamin D deficiency can largely be attributed to a reduction of intestinal
binding protein and lack of calcium recovered from feed (McCarthy et al., 1984). During vitamin D deficiency, growing birds
develop hypocalcemia, which, in turn, stunts skeletal development through widened cartilage at epiphyses of long bones and
weakened shafts (Noff et al., 1982; Long et al., 1984). For some reason, an abnormal blackening of the feathers also occurs
with some pigmented chicks (Glazener and Briggs, 1948). Once the skeleton has assumed adult size, a vitaminosis D is
obvious only with hens in production. Egg production and egg weight decrease while the eggshell thins as bone reserves are
progressively depleted (Vohra et al., 1979).
Hens in production cyclically release estrogen from the ovary to maximize 1,25-dihydroxy-vitamin D production
concurrent with eggshell formation (Castillo et al., 1979). As a result, levels of calcium-binding protein in the uterus
(Navickis et al., 1979) and calcium in the medullary bone (Takahashi et al., 1983) are altered to facilitate eggshell formation.
Vitamin D nutriture of the hen also influences its content in egg yolk and the subsequent need for this vitamin by the chick
(Bethke et al., 1936; Griminger, 1966; Stevens and Blair, 1985).
Vitamin D removed from the yolk is metabolized by the embryo as it is by the adult, and 1,25-dihydroxy-vitamin D is
the dominant metabolite (Bishop and Norman, 1975). An additional activity for this metabolite is recovery of calcium from
the shell at the chorioalloic membrane to support skeletal mineralization prior to hatching (Narbaitz, 1987). The yolk sac
membrane also responds to 1,25-dihydroxy-vitamin D at the same time, and a portion of the calcium from the shell is
transferred into the yolk for later use upon hatching (Clark et al., 1989); however, one or more of the other metabolites must
also be present if complete embryonic development and emergence from the shell is to occur (Ameenuddin et al., 1982).
The very low content of vitamin D in feedstuffs is generally ignored in feed formulation, and the complete requirement
is satisfied by using concentrated premixes. Overuse of vitamin D concentrates can lead to a toxicity. High levels of 1,25dihydroxy-vitamin D occur with a toxicosis, along with hypercalcemia and soft tissue mineralization (Morrissey et al., 1977;
Ratkowski et al., 1982). Leg problems may arise with growing birds because of bone calcium loss (Cruickshank and Sim,
1987), but few obvious changes occur with hens other than a general depression in performance (Ameenuddin et al., 1986).
Toxic levels of vitamin D may be transferred into the egg to create similar problems for the embryo; however, the
hypercalcemia occurs from shell resorption, and bone mineralization is enhanced (Narbaitz and Fragiskos, 1984).
Vitamin D in feed may not be totally available to poultry. This vitamin is susceptible to destruction by oxidation and
significant losses may occur unless supplemental antioxidants are used (Fritz et al., 1942). Also, mycotoxins in feeds interfere
with the utilization of dietary vitamin D (Bird, 1978; Gedek et al., 1978; Kohler et al., 1978). Losses of vitamin D because of
oxidation and poor utilization may result in a deficiency of the vitamin even though initial dietary concentrations of vitamin
D substantially exceed known requirements.
Vitamin E
Vitamin E is composed of an array of tocopherols derived from plant sources that act as antioxidants within the animal.
Hydrophobic areas of tissues, particularly cell membranes, are the sites of action for vitamin E (Erin et al., 1984), whereas
selenium is a cofactor for complementary antioxidant activities in the aqueous portion (Xu and Diplock, 1983). Dietary
vitamin E is absorbed from the intestine with fat, and its dissemination follows depletion of lipoprotein contents from
circulation (Massey, 1984). In turn, tissue vitamin E content parallels feed vitamin E levels, and tissues receiving the highest
proportions are intestine, liver, fat depots, and muscle (Astrup, 1979).
The amount of vitamin E needed to avoid a deficiency largely depends on the adequacy of the accompanying selenium
and on circumstances presenting oxidative threats to the system. An inadequacy of both vitamin E and selenium leads to
exudative diathesis, which is a subdermal accumulation of viscous blue-green-colored exudate from endothelial failures in
portions of the vascular system (Scott, 1966a). Myopathies of the gizzard, heart, and, to a lesser extent, the skeletal muscles
are also apparent. Skeletal muscles, particularly the breast, become more myopathic when the sulfur amino acids are also
deficient. Exudative diathesis can be eliminated and most myopathies can be greatly relieved when selenium alone is
increased (Combs and Scott, 1974).
Vitamin E deficiency symptoms that do not benefit from increased selenium are encephalomalacia (Hassan et al., 1985)
and the susceptibility of red blood cells to hemolysis (Dobinska et al., 1982). Degeneration of the Perkinji layer of cells in the
cerebellum results in nervous symptoms typified as sudden prostration with toes and legs outstretched, toes flexed, and head
outstretched. High concentrations of dietary PUFA lead to
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SIGNS OF NUTRITIONAL DEFICIENCIES IN CHICKENS AND TURKEYS
52
increased contents in cell membranes and, in turn, the additional susceptibility to oxidative stress may enhance the
possibilities of encephalomalacia (Budowski and Crawford, 1986). Other stressors such as ozone in the environment (Bartov
et al., 1981) or peroxidized fat (Budowski et al., 1979) or medium-chain fatty acids (Ikumo, 1980) contained in the feed also
increase the possibility of a vitamin E deficiency.
Adult fowl are less susceptible to a vitamin E deficiency than are actively growing chicks, and the symptoms differ.
Males become infertile because sperm become incompetent (Friedrichsen et al., 1980). Reduced egg production and
hatchability occur when both vitamin E and selenium are deficient over a prolonged period with hens (Latshaw and Osman,
1974). Although supplemental selenium can completely overcome these problems, chicks from these eggs are particularly
susceptible to encephalomalacia (Bartov and Bornstein, 1980) and muscular dystrophy (Ewen and Jenkins, 1967).
Adding excessive vitamin E to feed can have adverse effects. Nockels et al. (1976) reported that feeding 8,000 IU/kg
reduced body weight gain and gave a waxy appearance to the feathers. Should either vitamin D or vitamin K be marginal
when high levels of vitamin E are being fed, then rachitic bones and blood clotting failures, respectively, may occur (March
et al., 1973; Murphy et al., 1981; Franchini et al., 1988). However, dietary excesses approximating 100 to 500 IU/kg of feed
are advantageous to the oxidative stability of broiler (Lin et al., 1989) and turkey (Sheldon, 1984) meat products.
Vitamin K
Vitamin K is used as a cofactor to synthesize γ-carboxyglutamic residues from glutamic acid in proteins located in the
liver and bone. The liver protein is involved in the synthesis of several blood clotting factors, including prothrombin clotting
of blood (Suttie, 1987), and the bone protein, osteocalcin, is implicated in calcification of bone matrix (Hauschka et al., 1989).
Although inadequate dietary vitamin K alters bone osteocalcin, symptoms associated with the skeletal system are not as
apparent as blood clotting problems (Scott, 1966b; Hauschka and Reid, 1978b). Hemorrhaging may occur subcutaneously,
intermuscularly, and internally and may lead to anemia and the appearance of hypoplastic bone marrow. A greatly extended
blood clotting time may result in death from exsanguination. Vitamin K adequacy is usually measured in terms of
prothrombin clotting time with decalcified plasma (Griminger et al., 1970).
Dietary vitamin K may be of three sources. Vitamin K1, or phylloquinone, largely occurs in the leafy parts of plants.
Vitamin K2, or menaquinone, is of bacterial origin, particularly those bacterial located in the large intestine. Vitamin K3, or
menadione, has been synthesized and does not occur in nature as such. Antivitamin K compounds, whether synthetic
(Lowenthal and MacFarlane, 1965) or natural (Griminger, 1987), act as anticoagulants. Menadione generally exhibits the
greatest vitamin K activity (Dua and Day, 1966), except when anticoagulants are given and the converse occurs (Griminger,
1965). Dietary anticoagulants lead to vitamin K deficiency symptoms commensurate with the extent of toxicity (Veltmann et
al., 1981; Bai and Krishnakumari, 1986).
Inadequate vitamin K under practical circumstances is most likely to occur during the starting period, and
supplementation of the feed at this time is advantageous (Fritz, 1969). Starting feeds seldom contain forage meals, and a
poorly developed intestinal microflora together with the use of antimicrobials further reduces access to the vitamin (Bornstein
and Samberg, 1954). Nelson and Norris (1961a) showed that the inclusion of 0.1 percent sulfaquinoxaline increased the
chick's need for supplemental vitamin K by fourfold to sevenfold.
Adults usually have a well-developed intestinal microflora, and vitamin K inadequacies are unusual. Vitamin K2 is not
readily absorbed from the large intestine but it is digested after coprophagy of cecal excreta (Berdanier and Griminger, 1968).
The caging of hens minimizes coprophagy, and minimal amounts of vitamin K reach the egg (Cravens et al., 1941).
Griminger and Brubacher (1966) observed that dietary vitamin K3 is transferred to the yolk as vitamin K2, but vitamin K1 is
best transferred and remains as such.
Use of vitamin K by embryos parallels that by adults. A deficiency with the embryo alters bone metabolism, but no
physical deformities occur (Hauschka and Reid, 1978a). Adverse effects on blood clotting are not apparent until after
hatching, when hemorrhaging and mortality occur should trauma be encountered (Griminger, 1964).
Thiamin (Vitamin B1)
Thiamin is a cofactor for several enzymes catalyzing decarboxylationand transketolation-type reactions. Although the
activity of all these enzymes is depressed in a thiamin deficiency, the accrual of pyruvic acid from decreased brain pyruvic
oxidase seems to manifest the most symptoms (Lofland et al., 1963). Ataxia and awkward backward flexions of the head and
neck are typical nervous symptoms (Gries and Scott, 1972b). Deficient birds can rapidly detect and discriminate against feeds
that do not provide the vitamin (Hughes and Wood-Gush, 1971) and are high in carbohydrate content (Thornton and Shutze,
1960).
Most complete feeds satisfy the thiamin requirement because grains and their by-products usually contain adequate
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53
amounts. Thiamin is unstable to heat at neutral and alkaline pH (Dwivedi and Arnold, 1973), and pelleting (Guo and
Summers, 1969) or extrusion (Beetner et al., 1974) under these circumstances facilitates loss. Amaranth is very low in
thiamin, and the level is reduced further if it is heated to destroy growth-inhibiting properties (Laovoravit et al., 1986).
Inclusion of certain fish meals having enzymes capable of destroying thiamin may also decrease dietary content (Ishihara et
al., 1974; Bryan et al., 1975). Use of medicants acting as a thiamin antagonist can also cause a deficiency (Ott et al., 1965;
Shindo et al., 1972).
The hen transfers thiamin to the egg in proportion to dietary content (Polin et al., 1963). Although the dietary
inadequacies possible under practical terms do not affect breeder flock productivity, high mortality of embryos occurs prior to
hatching and chicks that hatch express a polyneuritis (Polin et al., 1962; Charles et al., 1972).
Riboflavin (Vitamin B2)
Riboflavin acts as a cofactor for many enzymes involved in oxidation-reduction. Erythrocyte glutathione reductase (Lee,
1982) and liver xanthine dehydrogenase (Chou, 1971) are two enzymes in fowl shown to need riboflavin, and their activities
reflect dietary adequacy. Prior to the development of concentrated riboflavin sources, milk products were incorporated in feed
to avoid deficiencies (Culton and Bird, 1940).
Riboflavin deficiencies lead to neurological problems, particularly with the sciatic and brachial nerves, where myelin
degeneration, Schwann cell proliferation, and axis cylinder fragmentation have been observed (Phillips and Engel, 1938).
Symptoms involving the legs of chickens appear as splay and hock resting postures, and curling of the toes occurs to a lesser
extent (Wyatt et al., 1973a; Ruiz and Harms, 1988a). Turkey poults (Ruiz and Harms, 1989a) and pheasants (Scott et al.,
1959) exhibit similar symptoms as the chick, whereas ducks (Fritz et al., 1939) and geese (Serafin, 1981) are more likely to
have a bowing of the legs in conjunction with perosis. Goff et al. (1953) noted that increased hematocrit, increased mean
corpuscular volume, decreased mean hemoglobin concentration, and a marked heterophil leucocytosis appeared in the chick
prior to neurological manifestations.
Adult cockerels can endure a riboflavin-deficient feed for a prolonged period before neurological and blood problems
similar to those of the growing chick appear (Arscott, 1972). Deficiency symptoms can be reversed upon riboflavin
administration to adults, but correction with growing birds becomes increasingly difficult as expression progresses.
Laying hens transfer riboflavin into the yolk and albumen by hormonally induced binding proteins in the liver and
oviduct, respectively (Hamazume et al., 1984). Saturation of these carriers is dependent on dietary riboflavin content (White
et al., 1986), and an inadequacy is more likely to adversely affect embryonic development than harm the hen (Tarhay et al.,
1975). Severe inadequacies cause death of embryos at 60 hours incubation because of circulatory system failures (Romanoff
and Bauernfeind, 1942). Moderate inadequacies result in deaths at 14 days incubation, with the appearance of shortened
limbs, malformed mandibles, and clubbing of the down. Marginal deficiencies further delay mortality until pipping, and
symptoms are largely dwarfism with clubbed down.
Niacin
Niacin represents nicotinic acid and nicotinamide, both of which have similar activity in fowl (Ruiz and Harms, 1988b).
Many enzymes in glycolysis, lipogenesis, and energy metabolism use niacin as a cofactor. Tryptophan may be converted into
niacin; however, the efficiency is poor and not recommended as a substitute for diet supplementation (Ruiz and Harms, 1990).
Availability of niacin in grain and grain by-products is generally low (Manoukas et al., 1968; Yen et al., 1977); thus
their contribution in determining dietary adequacy is usually ignored. Chicks at hatch have considerable tryptophan contained
in the protein of the yolk; thus a niacin deficiency will not readily occur unless the feed is low for both the amino acid and the
vitamin (Snell and Quarles, 1941). Briggs et al. (1943) reported that 2 weeks were required to provoke a deficiency with
chicks and that an inflammation of the oral cavity and occasional poor feathering, dermatitis, and perosis—a malformation of
the bones—were the primary symptoms. Turkey poults (Ruiz and Harms, 1988b), pheasants (Scott et al., 1959), ducks
(Heuser and Scott, 1953), and goslings (Serafin, 1981) all expressed perosis as the primary deficiency symptom.
Biotin
Biotin acts as a cofactor for enzymes performing carboxylations. Acetyl coenzyme A carboxylase, which participates in
fatty acid synthesis, and pyruvate carboxylase, which enables gluconeogenesis from intermediates in the Kreb's cycle, are
both affected by biotin nutriture (Whitehead and Bannister, 1980; Watkins and Rogel, 1989). Biotin tends to concentrate in
liver, kidney, and bone, the primary sites of activity of enzymes requiring this vitamin (Frigg and Torhorst, 1982). Analysis
of complete feeds indicates that adequate biotin is
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SIGNS OF NUTRITIONAL DEFICIENCIES IN CHICKENS AND TURKEYS
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present; however, low availability of biotin from certain grains may result in marginal concentrations in comparison with
biotin requirements (Frigg, 1976).
Symptoms of a biotin deficiency are skin lesions appearing on the foot pad, shank, and toes, together with eye
exfoliation and exudative dermatitis (Marusich et al., 1970). Skin lesions can be related to alterations in the fatty acid
composition of associated waxes (Logani et al., 1977). Low dietary fat and the necessity for fatty acid synthesis lead to an
abnormal array of fatty acids that predisposes poultry to a fatty liver and kidney syndrome (FLKS) (Whitehead and Randall,
1982). Subjecting these birds to a fast such that gluconeogenesis is accelerated precipitates a high death rate from lack of
glucose (Whitehead and Siller, 1983). Tibiotarsal bones are frequently longitudinally distorted. Presumably, reduced biotin
prevents ready formation of prostaglandins from essential fatty acids, and bone growth fails to respond to stresses during
development (Watkins et al., 1989).
Biotin-binding proteins are found in the yolk and albumen of eggs (Bush et al., 1988). The amount of biotin associated
with the yolk binding protein changes with biotin content in the feed. Hatchability is affected when the feed is deficient
(White et al., 1987). Embryonic mortality because of inadequate biotin occurs largely during the last 3 days of incubation.
Dwarfing, chondystrophy, and deformities of the mandibles and skeleton appear at that time (Couch et al., 1947).
Chicks hatched from breeder hens given marginal dietary biotin have increased risk of a deficiency (Whitehead et al.,
1985). Provoking a deficiency is dependent on many factors, particularly those affecting supplementary biotin synthesis by
microbes in the ceca and coprophagy. Caging and use of probiotics and medicants in the feed are influential in this respect
(Leeson, 1982).
Pantothenic Acid
Pantothenic acid serves as a prosthetic group with coenzyme A and thereby is essential in energy metabolism.
Inadequate pantothenic acid not only reduces the productive use of available energy (Beagle and Begin, 1976; Cupo and
Donaldson, 1986) but also impairs detoxification mechanisms that depend upon acetylation (Kietzmann, 1981). Grains
contain low concentrations of pantothenic acid, and complete feeds are usually marginal in satisfying the requirement
(Southern and Baker, 1981; Ruiz and Harms, 1989b).
Deficiency symptoms are associated with the skin and nervous system of growing chicks (Gries and Scott, 1972b). Skin
lesions include crusts and scabs, which first appear at the angles of the eyes and beak. Lesions on the feet are seldom and
slight. Biotin deficiency symptoms are similar except lesions on the feet are more severe and appear before those on the head.
Although an extensive ataxia also occurs, lesions associated with the nervous system are difficult to detect. Turkey poults
present the same symptoms as chicks (Kratzer and Williams, 1948a), but poor feathering is the most prevalent deficiency sign
in pheasants and quail (Scott et al., 1964).
Adult cockerels receiving inadequate pantothenic acid have reduced semen volume and fertility as well as skin lesions
(Goeger and Arscott, 1984). Considerably higher levels of pantothenic acid are needed by chicken and turkey hens to
maintain hatchability than for egg production (Kratzer et al., 1955; Balloun and Phillips, 1957a). Embryonic mortality occurs
from about 14 days incubation or thereafter, depending on the extent of pantothenic acid inadequacy (Beer et al., 1963).
Chicks that hatch are of poor quality and have variable degrees of subcutaneous hemorrhaging and edema (''stunted chick
disease").
Pyridoxine (Vitamin B6)
Pyridoxine, pyridoxal, and pyridoxamine are the 3 active forms of vitamin B6. Vitamin B6 is a cofactor in
decarboxylation and transamination reactions of amino acids. Decarboxylations lead to at least four amines that affect
nervous system functioning. Transaminations of certain glycolysis and Kreb's cycle intermediates form most of the
nonessential amino acids, whereas the reverse is the basis of gluconeogenesis from protein. Aspartic transaminase in the liver
(Lee et al., 1976) and plasma glycine-serine ratio (Sifri et al., 1972) have been employed to evaluate vitamin B6 nutriture.
The vitamin B6 content of complete feeds usually satisfies most requirements (Scheiner and DeRitter, 1968). However,
the vitamin availability is dependent on the digestibility of each feedstuff (Heard and Annison, 1986). The dietary
requirement level may increase as dietary protein increases (Daghir and Shah, 1973), or due to the presence of linatin when
linseed meal is used (Kratzer and Williams, 1948b; Klosterman et al., 1967). The inclusion of certain drugs that act as
competitive inhibitors may also increase the dietary requirement (Fuller and Dunahoo, 1959).
Symptoms exhibited by vitamin-B6-deficient chicks differ with the extent of the inadequacy (Daghir and Balloun, 1963;
Gries and Scott, 1972a). A severe deficiency produces an ataxia in combination with nervousness and intermittent episodes of
hyperactivity. Prominent pathological findings include hemorrhages at various locations, particularly primary wing feather
follicles, and gizzard erosions. Marginal vitamin B6 deficiencies are most likely to be expressed as a perosis because of
problems with bone growth. Miller (1963) observed high proportions of pendulous crops with vitamin-B6-deficient chicks.
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Blood alterations are also typical of a vitamin B6 inadequacy. An extreme deficiency leads to a microcytic,
polychromatic hypochromic anemia in conjunction with atrophy of the spleen, thymus, and bursa of Fabricius (Asmar et al.,
1968). Marginal deficiencies provoke a microcytic, normochromic polycythemia (Blalock and Thaxton, 1984), and deficient
chicks show a decreased immunoglobulin M and immunoglobulin G response to antibody challenge (Blalock et al., 1984).
Although specific symptoms of vitamin B6 deficiency are not obvious in adult chickens, deficient hens lose body weight
and exhibit reduced egg production (Attar et al., 1967). Deficient hens also have relatively low serum glutamic-oxaloacetic
acid transaminase activities and high serum nonprotein nitrogen levels (Attar et al., 1967). The vitamin B6 content of eggs
reflects that in the feed, and the level necessary to maintain egg production is one-half of that required for hatchability (Fuller
et al., 1961). Characteristics of vitamin-B6-deficient embryos have not been reported, but antivitamins injected into eggs
cause early deaths (Landauer, 1967).
Folic acid
Folacin represents folic acid (pteroyl-γ-monoglutamic acid) and the array of extended glutamic acid conjugates.
Enzymes engaged in one-carbon metabolism use folic acid as a cofactor in methyl and methylene group synthesis. Dietary
folacin is absorbed and converted to the reduced form (5-methyl-tetrahydrofolic acid) by the intestine and is distributed
throughout the body.
Although most complete feeds provide sufficient folic acid from their natural ingredients, marginal inadequacies are
possible (Cropper and Scott, 1967). The requirement decreases with age because diminished growth rate reduces the need for
deoxyribonucleic acid synthesis (Naber et al., 1957; Balek and Morse, 1976). Accentuated formation of uric acid with
excessive dietary protein increases the folic acid requirement (Creek and Vasaitis, 1963), as does inadequate choline (Young
et al., 1955) and serine (Rabbani et al., 1973). Use of medicants that antagonize folic acid formation by cecal microflora and
management that prevents coprophagy also increases the dietary requirement (Stokstad and Jukes, 1987).
The most obvious symptom of inadequate folic acid is perosis with the chick (Daniel et al., 1946) and cervical paralysis
with turkey poults (Miller and Balloun, 1967). Macrocytic anemia, abnormal nuclear bodies in erythrocytes, and numerous
mitoses and hypersegmented granulocytes occur with marginal deficiencies when no physical symptoms are manifested
(Maxwell et al., 1988).
Inadequate folic acid with the hen impairs the oviduct's response to estrogen and ability to form albumen (Anderson and
Jackson, 1975; Burns and Jackson, 1979). More folic acid is needed to sustain hatchability than egg production; thus the
embryo will suffer before the hen (Sunde et al., 1950a). High embryonic mortality occurs around 20 days of incubation, and
the dead from severely depleted hens exhibit a marked bending of the tibiotarsus, and, to a lesser extent, syndactyly and
deformed mandibles. Chicks that successfully emerge are stunted and have feathers that are poorly developed and abnormally
pigmented (Lillie et al., 1950).
Vitamin B12 (Cobalamin)
Vitamin B12 is a cofactor for enzymes transferring one-carbon units and catalyzing rearrangements in the carbon
skeleton of several metabolic intermediates. In fowl, vitamin-B12-mediated one-carbon transfers involve methionine, serine,
choline, and thymidine (Gillis and Norris, 1949; Henderson and Henderson, 1966; Langer and Kratzer, 1967), whereas the
interconversion of methylmalonyl coenzyme A to succinyl coenzyme A is one of the rearrangement reactions requiring
vitamin B12 (Ward et al., 1988).
The spleen, bone marrow, liver, kidney, and skin have high concentrations of vitamin B12 (Monroe et al., 1952).
Although plant feedstuffs are devoid of vitamin B12, its availability from animal products and cecal microflora after
coprophagy makes deficiencies unlikely (Milligan et al., 1952). Deficiencies in chicks have been created by greatly
increasing dietary protein content such that carbon rearrangement enzyme activities are accentuated (Rys and Koreleski,
1974; Patel and McGinnis, 1980; Ward et al., 1985). Poor feathering and mortality are the most obvious symptoms of a
vitamin B12 deficiency, and gizzard erosions may also appear (Mushett and Ott, 1949; Milligan et al., 1952).
Yacowitz et al. (1952) fed a high-protein all-vegetable diet devoid of vitamin B12 to hens in cages and reported a
reduction in hatchability. Olcese et al. (1950) observed that most embryonic mortality due to vitamin B12 deficiency in hens
occurs at about 17 days of incubation, with atrophy of the leg musculature and hemorrhaging common. Ferguson et al. (1955)
further observed fatty organs, dwarfing, and edema.
Choline
Choline may be synthesized in fowl; however, the extent is limited, and supplementation is necessary when demand
exceeds biosynthesis capacity. Choline serves a diversity of needs, particularly as a component of phospholipids for the
formation of membranes and lipoproteins. Choline also acts as a methyl donor, and its use in this respect becomes important
when de novo synthesis of one-carbon units cannot meet demand.
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SIGNS OF NUTRITIONAL DEFICIENCIES IN CHICKENS AND TURKEYS
56
Need for supplemental choline is the greatest with the starting bird because all facets of use are likely to be maximal
(Seifter et al., 1972; Pesti et al., 1980). As growth diminishes, the necessity for choline supplementation disappears (Molitoris
and Baker, 1976). Perosis is the primary symptom of a choline deficiency in chicks (Fritz et al., 1967) and turkey poults
(Evans et al., 1943), whereas Bobwhite quail develop enlarged hocks and bowed legs (Serafin, 1974).
Estrogenic hormones greatly accentuate the choline need for phospholipid synthesis in the hen's liver to support yolk
formation (Vigo and Vance, 1981). Supplemental choline may relieve the hepatic accumulation of fat and improve egg yolk
formation (Schexnailder and Griffith, 1973; Tsigabe et al., 1988). Minimal dietary choline does not affect hatchability with
either chickens (Gish et al., 1949) or turkeys (Ferguson et al., 1975), but Japanese quail and their developing embryos readily
express general signs of deficiency (Latshaw and Jensen, 1971, 1972).
MINERAL DEFICIENCIES
Calcium and Phosphorus
Bone formation is highly dependent on the dietary concentrations of calcium and phosphorus as well as on adequate
intake of vitamin D3 (Hart et al., 1922; Dunn, 1924; McGowan and Emslie, 1934). Deficiency of any one of these nutrients
will result in rickets. Poor growth may also be a sign of calcium or phosphorus deficiency.
Dietary excesses of either calcium or phosphorus should be avoided because such excesses can hinder the intestinal
absorption of other mineral elements (Gutowska and Parkhurst, 1942; Schaible and Bandemer, 1942; Migicovsky and Emslie,
1947). The phosphorus that comes from plant products (that is, phytin) should not be depended on to fulfill the phosphorus
requirement for two reasons: it is not readily available in its natural form to the bird, and it may bind calcium, zinc, iron, and
manganese so as to render them unavailable (Nelson and Walker, 1964; Kratzer and Vohra, 1986).
Pullets at the beginning of the laying period undergo considerable metabolic stress associated with adjustment to the
need to supply approximately 2.4 g of calcium daily to the oviduct for shell formation (Mueller et al., 1964; Hurwitz and Bar,
1971; Scott et al., 1971). Some birds mobilize large amounts of calcium from their skeleton during this period, and the bones
may become so demineralized that the birds are unable to stand and appear paralyzed. The sternum and rib bones are
frequently deformed, and all bones are easily broken. Dietary management to prevent this condition (generally termed "cagelayer fatigue" but more precisely described as osteoporosis) has not been devised (Roland et al., 1968).
Magnesium
When fed a diet very deficient in magnesium, chicks grow slowly for about 1 week and then stop growing and become
lethargic. Chicks fed diets marginal in magnesium may grow quite well but exhibit reduced levels of plasma magnesium and
symptoms of neuromuscular hyperirritability when disturbed (Almquist, 1942; Bird, 1949). Chicks show a brief convulsion
and then enter a comatose state from which they usually recover, but sometimes death occurs.
A magnesium deficiency in laying hens results in a rapid decline in blood magnesium level, withdrawal of magnesium
from bone, decline in egg production, and, eventually, a comatose state and death (Cox and Sell, 1967). Magnesium content
and hatchability of eggs also are reduced when hens are fed magnesium-deficient diets (Sell et al., 1967; Hajj and Sell, 1969).
Increasing either the calcium or the phosphorus content of the diet accentuates magnesium deficiency (Nugara and Edwards,
1963). Normally, adequate magnesium is present in the natural ingredients of practical diets to meet the requirements of
poultry.
Manganese
Manganese deficiency in chicks and poults results in perosis or slipped tendon (Wilgus et al., 1937; Ringrose et al.,
1939). Deficiencies of other nutrients, such as choline and biotin, may also be involved in inducing perosis (Jukes, 1940;
Jukes and Bird, 1942). The usual signs of perosis are swelling and flattening of the hock joint, with subsequent slipping of the
Achilles tendon from its condyles. The tibia and the tarsometatarsus may exhibit bending near the hock joint and lateral
rotation. One or both legs may be affected. A shortening and thickening of the long bones of the wings and legs are also
observed. The disorder, insofar as manganese is concerned, is aggravated by excess dietary calcium and phosphorus
(Schaible and Bandemer, 1942).
In laying and breeding birds, manganese deficiency results in lowered egg production, reduced eggshell strength, poor
hatchability, and reduced fertility. Manganese-deficient embryos exhibit shortening of the long bones, parrot beak, and wiry
down (Lyons and Insko, 1937; Caskey et al., 1939).
Potassium, Sodium, and Chlorine
A deficiency of potassium results in high mortality and retarded growth of chicks and causes reduced egg
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SIGNS OF NUTRITIONAL DEFICIENCIES IN CHICKENS AND TURKEYS
57
production and eggshell thickness in laying hens (Ben-Dor, 1941; Gillis, 1948; Leach, 1974). It is not usually necessary to
add potassium to practical feed formulations, since such formulas generally contain about 0.7 to 1.0 percent potassium.
A deficiency of sodium in chicken diets results in poor growth, increased adrenal weight, and decreased egg production
(Burns et al., 1952, 1953; Nott and Combs, 1969). Frequently, sodium supplementation is minimized to reduce the moisture
level in the excreta.
Signs of chlorine deficiency in chicks include poor growth, mortality, hemoconcentration, and reduced blood chlorine
level (Leach and Nesheim, 1963). Chlorine-deficient chicks show a nervous condition resembling tetany and fall forward
with legs extended backward when stimulated by a sharp noise.
Iodine
Iodine is necessary for the synthesis of thyroid hormones. Iodine deficiency results in goiter, which is the enlargement of
the thyroid glands (Wilgus et al., 1953; Rogler et al., 1959a). The glands may increase to many times their usual size. If the
deficiency is not too severe, the increased efficiency of the enlarged gland in "trapping" iodine from the bloodstream may
compensate for the low dietary concentration. When this is the case, the production of thyroid hormones is normal, although
the thyroid glands are enlarged.
Inadequate production of thyroid hormones results in poor growth, egg production, and egg size. Iodine deficiency in
breeders results in low iodine content of the egg and, consequently, decreased hatchability and thyroid enlargement in the
embryos.
Copper
Copper deficiency in poultry causes an anemia in which the red blood cells are small and low in hemoglobin (Elvehjem
and Hart, 1929). Bone deformities can occur (O'Dell et al., 1961). Pigmentation of feathers in New Hampshire and Rhode
Island Red chickens is reduced (Hill and Matrone, 1961). Copper is required for the activity of the enzyme needed for the
cross-linking of lysine in the protein elastin (O'Dell et al., 1961; Starcher et al., 1964). Dissecting aneurism of the aorta
occurs in birds deficient in copper because of the defect in elastin formation. Copper deficiency also results in marked cardiac
hypertrophy (Carlton and Henderson, 1963).
Iron
Iron deficiency in chickens and turkeys causes an anemia in which the red blood cells are reduced in size and low in
hemoglobin (Elvehjem and Hart, 1929). In red-feathered chickens, pigmentation does not occur when the diet is deficient in
iron (Hill and Matrone, 1961; Davis et al., 1962).
Selenium
Selenium is closely associated with vitamin E and other antioxidants in practical feed formulation. The principal sign of
deficiency in chicks is exudative diathesis (Creech et al., 1957; Patterson et al., 1957; Nesheim and Scott, 1958). A
requirement for selenium supplementation, even in the presence of vitamin E, is demonstrated by the poor growth, muscular
dystrophy, and mortality of chicks fed purified diets or diets based on grains produced on low-selenium soils (Nesheim and
Scott, 1958). Selenium is required for prevention of myopathies of the gizzard and heart in turkeys (Walter and Jensen, 1963;
Scott et al., 1967). Pancreatic fibrosis, with resultant reductions in the pancreatic output of lipase, trypsinogen, and
chymotrypsinogen, has also been associated with selenium deficiency (Thompson and Scott, 1970; Gries and Scott, 1972c).
Selenium is a structural component of glutathione peroxidase, an enzyme needed to quench peroxides generated during
metabolism (Rotruck et al., 1973).
There is wide variability in the amount and availability of selenium in the soils of different geographic areas (Scott and
Thompson, 1971; Scott, 1973). Consequently, cereals and plant-derived feedstuffs are variable sources of selenium. Grains
from some areas contain sufficient selenium to render them toxic to chicks. The effects of toxic levels of selenium are listed
in Table 8-1. The amount of supplementary selenium permissible in diets is regulated in the United States and Canada.
Zinc
Zinc has many biochemical functions. Deficiency causes retarded growth and frayed feathers (O'Dell et al., 1958;
Sullivan, 1961). The extent of fraying varies from almost no feathers on the wings and tail to only slight defects in the
development of some of the barbules and barbicels. The long bones of the legs and wings are shorter and thicker than normal
(Kratzer et al., 1958; Morrison and Sarett, 1958; O'Dell et al., 1958). The hock joint may be enlarged. Layer and breeder diets
deficient in zinc reduce egg production and hatchability (Kienholz et al., 1961).
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TOXICITY OF CERTAIN INORGANIC ELEMENTS
58
8
Toxicity of Certain Inorganic Elements
Current information on toxic dietary levels of inorganic elements for poultry is summarized in Table 8-1. A similar
summary that describes the mineral tolerances of animals has been provided by the National Research Council (1980b).
Toxicity, as defined here, is any adverse effect on performance. Reduced growth rate is the most common criterion used to
indicate the specific level at which a particular mineral is toxic. Although most of the information in the table was obtained
from experiments in which the mineral was added in the form of an inorganic compound, organic compounds served as the
source of minerals in some reports. For instance, some of the information on the toxicity of selenium was obtained by feeding
seleniferous wheat.
The toxicity of a mineral is influenced by the nature of the compound in which it is present (for example, methyl
mercury is much more toxic than mercuric chloride). Toxicity may also be influenced markedly by the composition of the
diet, particularly with respect to other minerals and chelating agents. Selenium included in the diet at 10 ppm reduces the
growth rate, but when it is fed in combination with 1,000 ppm of silver, a level as high as 40 ppm does not reduce growth
(Jensen, 1975a). Copper at a level of 800 ppm in a practical turkey diet is not toxic, but 50 ppm of copper in a purified diet
reduces growth. The toxicity of copper is modified by the sulfur amino acid content of the diet. Vanadium is much more toxic
in a purified diet than in a practical diet, and the toxicity is increased by adding lactose to the practical diet (Hafez and
Kratzer, 1976). Conversely, vanadium toxicity is reduced by including cottonseed meal in the diet (Berg, 1965; Berg and
Lawrence, 1971; Sell et al., 1986a). In many instances, a high dietary level of one mineral antagonizes another element,
resulting in a physiological deficiency of minerals essential for the animal. Because many different factors affect the quantity
of a mineral needed to produce toxicity, diverse observations have been reported on the toxic effects of any given mineral.
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TOXICITY OF CERTAIN INORGANIC ELEMENTS
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COMPOSITION OF FEEDSTUFFS USED IN POULTRY DIETS
61
9
Composition of Feedstuffs Used in Poultry Diets
Feed formulation involves the judicious use of feed ingredients to supply in adequate amounts and proportions the
nutrients required by poultry. Because it is impractical to analyze each batch of feedstuff for its nutrient content, reliance
must be placed on feedstuff composition data that have been compiled on the basis of many laboratory analyses. Feedstuffs
vary in composition. The nutrient values given in the following tables are averages reflecting the concentrations of nutrients
most likely to be present in the feedstuffs commonly used in poultry feeds.
Feedstuff composition data presented in this edition (Tables 9-1 and 9-2) were obtained from several sources, including
the United States-Canadian Tables of Feed Composition (National Research Council, 1982), the Association of American
Feed Control Officials, commercial firms, and individual scientists. In many instances, the values have been changed to
reflect results of analyses of feed ingredients obtained from contemporary crop cultivars and recently employed processing
methods. Additional information provided in the composition tables include nitrogen-corrected true metabolizable energy
(TMEn) data for many feed ingredients and information on the true digestibility of amino acids for numerous feedstuffs. Also,
equations are provided to estimate the amino acid concentration of certain ingredients on the basis of proximate analysis or
on the basis of the protein content of the ingredients.
From a nutritional point of view, there is no "best" diet formula in terms of ingredients that are used. Ingredients should,
therefore, be selected on the basis of availability, price, and the quality of the nutrients they contain. Certain ingredients
invariably constitute the greatest part of diets, in terms of both amount and cost. Cereal grains and fats are the primary energysupplying ingredients, and oilseed meals and animal-protein meals are used commonly as major sources of amino acids.
Some important nutritional characteristics of many energy- and protein-supplying ingredients are discussed in this chapter.
Sulphur, which are common contaminants in feedstuffs, and their effects are discussed in the final section.
CEREAL GRAINS
Bushel weights (bulk densities) of cereal grains are used in commerce to establish market grades and prices. Bushel
weights of grains also have been used as criteria of feeding value, and in some instances this practice seems justified for
poultry. For example, at standard moisture levels there is a strong relationship between bushel weight and general feeding
value of oats and barley. An increase in bushel weight of these grains is a reflection of an increase in the proportion of the
meaty kernel and a decrease in the proportion of fibrous hull. Thus there is a definite increase in the metabolizable energy (ME)—
and usually protein—content of barley and oats as bushel weight increases. Similarly, there seems to be a direct relationship
between the ME content of grain sorghum and wheat as bushel weight increases over a wide range. A relationship between
bushel weight and the ME content of corn is not so evident. In situations in which corn, sorghum, or wheat fails to achieve
maturity because of early frost or early harvest, there usually are decreases in the starchy endosperm portion of the grain and
bushel weight and ME content are usually low. Regression equations relating the ME of corn to various factors such as
moisture content at harvest and bushel weight have been reported (Leeson and Summers, 1975, 1976b; Leeson et al., 1977b).
Ranges in bushel weight that may be encountered with different grains are shown in Table 9-3.
The feeding value of grain sorghums (milo) is markedly
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COMPOSITION OF FEEDSTUFFS USED IN POULTRY DIETS
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COMPOSITION OF FEEDSTUFFS USED IN POULTRY DIETS
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COMPOSITION OF FEEDSTUFFS USED IN POULTRY DIETS
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influenced by the tannin content of the grain. Development of high-tannin or ''bird-resistant" varieties has allowed
increased production of sorghum in areas where bird predation had previously limited yields; however, the presence of
tannins in these cultivars may reduce their nutritional value. Tannins cause a binding and precipitation of dietary proteins and
digestive enzymes (Butler et al., 1984) and may reduce both the amino acid (Armstrong et al., 1974) and the energy
digestibility
TABLE 9-3 Ranges in Weights per Unit of Volume for Selected Feedstuffs at Standard Moisture
Feedstuffs
Pounds per Bushel
Kilograms per Hectoliter
Barley
36–48
45–62
Corn
46–56
59–72
Oats
22–40
28–52
Sorghum (milo)
51–57
66–74
Soybeans
49–56
63–72
45–63
58–81
Wheat
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Moisture (%)
16.0
15.5
16.0
15.5
13.0
15.5
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(Gous et al., 1982) of the diet. The ME of grain sorghums can be predicted from their tannin content by the following
equation (Gous et al., 1982):
Although wheat was once considered too expensive for use in animal feeds, increased production in recent years has
resulted in more extensive use in poultry diets. In general, wheat has about 90 percent of the ME value of corn. The protein
and amino acid composition varies widely and is influenced by genetic and environmental factors. Most wheat varieties have
been developed for various baking properties, although some breeders have developed varieties designed primarily for animal
feeds (Bowyer and Waldroup, 1987). The nutrient sources in wheat are easily digested (McNab and Shannon, 1974). Feeding
trials with broilers, layers, and turkeys indicate that wheat can be effectively used to provide a major portion of the energy in
these diets (Waldroup et al., 1967; Lillie and Denton, 1968; Petersen, 1969). But because wheat has no carotenoid pigments,
adjustment is made when skin or yolk pigment must be maintained.
One vitamin that must be considered with wheat feeding is biotin. Although the total biotin content in wheat exceeds
that in corn, the biological availability in wheat is low (Frigg, 1976). A condition known as fatty liver and kidney syndrome
(FLKS) has frequently been observed in all species of poultry when wheat is used extensively. Biotin supplementation should
be considered when wheat provides more than 50 percent of the cereal grain.
Notwithstanding differences in bushel weight, the protein content of grains (dry matter basis) often varies a great deal
from batch to batch. This variation may be the result of genetic constitution, soil fertility, time of harvest, and other factors.
The protein concentration of grains can be determined readily for feed formulation purposes. It should be recognized,
however, that the amino acid composition of protein in a specific grain does not remain constant as protein concentration
changes. In some instances, the concentrations of essential amino acids in protein increase, but, in other instances, they
decrease. For example, there is a marked inverse relationship between the protein content of wheat or sorghum grain and the
lysine concentration in the protein. As protein content increases, lysine in the protein decreases. This relationship is most
prominent within cultivars of wheat and sorghum grains and is the result of a shift among the major proteins within these
grains, whereby the proportion of prolamine (low in lysine) increases at the expense of other proteins high in lysine. Certain
other amino acids (such as arginine, methionine, and cystine) may be affected similarly. An inverse relationship between
protein content and concentration of certain essential amino acids in the protein also has been reported for cultivars of barley,
corn, oats, and rice. The alterations in amino acid composition with increasing protein concentration generally are less with
these grains than with wheat and milo.
Recently, much research has been focused on the selection of cultivars of grains in which the concentrations of both
protein and selected amino acids within the protein may be increased. Examples include high-lysine corn and high-protein
barley. The quantities of these grains available for feeding to poultry are limited at the present time.
PROTEIN SUPPLEMENTS
A number of the feedstuffs used to supply supplementary protein to poultry diets may contain naturally occurring toxic
or potentially toxic compounds. In many instances, the nutritive value of the protein supplement can be markedly influenced
by the method used in processing the protein supplement.
Cottonseed Meal
Cottonseed meal, for example, may contain gossypol pigments. Free gossypol forms complexes with iron in the feed,
intestinal tract, blood, and egg yolk, leading to possible iron deficiency or to discoloration of the yolk. Under extreme heat
during processing, the gossypol may also form complexes with lysine, severely reducing the digestibility. The amount of
gossypol present in cottonseed meal is variable and depends on the cultivar and the manufacturing procedures. In general,
meals produced by the prepress solvent method are lowest in free gossypol, have greater lysine digestibility, and are the
preferred meal for poultry (Phelps, 1966). Gossypol adversely affects the bird, with younger birds being less tolerant than
older birds. Hens consuming gossypol may lay eggs with olive-discolored yolks, with the incidence related to the amount of
free gossypol consumed. The discoloration may be evident in the newly laid egg, but it more often becomes apparent after
storage. Addition of soluble iron salts to bind the free gossypol may enable the use of cottonseed meals, where this is
economically feasible (Waldroup, 1981). The presence of cyclopropenoid fatty acids and gossypol in cottonseed meals and
oil may also cause a pinkish color in the egg whites.
Rapeseed Meals
Rapeseed meals manufactured from many varieties of rapeseed contain goitrogenic, or progoitrogenic, compounds
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70
(glucosinolates) at sufficiently high concentrations to reduce growth rate and egg production when fed to poultry. Canadian
plant geneticists have been successful in developing rapeseed cultivars, called canola, that contain negligible quantities of
glucosinolates in the seed. Meals manufactured from these cultivars are called canola meal.
Inclusion of rapeseed meals in the diet of brown-egg layers sometimes results in the production of eggs with a "fishy" or
off-flavor taint. This taint is due to the presence of excess amounts of trimethylamine (TMA) in the yolk. Deposition of TMA
in yolks by certain strains of chickens is due to the presence of an autosomal semidominant gene that has variable expression
depending upon various environmental factors including the inclusion rate of rapeseed meal. Although some brown-egg
strains carry this trait, white-egg strains do not. This genetic defect reduces the synthesis of TMA oxidase enzyme, leading to
increased quantities of TMA in the metabolic pool. Rapeseed contains variable levels of sinapine, a potent inhibitor of TMA
oxidase. Low-glucosinolate cultivars have less drastic effects on egg taint but do not completely correct the situation.
Therefore care should be taken in feeding rapeseed or canola meals to hens that produce brown-shelled eggs.
Soybean Meal
Soybeans contain compounds that inhibit the activity of the proteolytic enzyme trypsin (Read and Haas, 1938). They
also contain other antinutrients, including hemagglutinins or lectins, which contribute to growth depression (Ham et al., 1945;
Chernick et al., 1948; Coates et al., 1970; Liener, 1980). Ingestion of the antitryptic substances induces enlargement of the
pancreas.
The trypsin inhibitor is inactivated by heat treatment of soybean meal. The heat treatment must be carefully controlled
because overheating can result in deterioration of protein quality. On the basis of the assumption that the urease enzyme in
raw soybeans is denatured at approximately the same rate as the trypsin inhibitor, and because it is easier to determine urease
activity than trypsin inhibitor, urease assays (Caskey and Knapp, 1944) have generally been used by the feed industry in
monitoring soybean meal quality. However, some studies indicate that there is not a direct relationship between the activities
of the two enzymes (Albrecht et al., 1966) and that the rates of destruction of urease and the trypsin inhibitor are not equal
under different processing conditions (McNaughton and Reece, 1980).
The feed industry in the United States has long used a maximum urease rise of 0.2 pH units as the standard for
processing soybean meal for all types of livestock feeds. However, studies show that meals with a urease value up to 0.50 pH
units are acceptable in poultry feeds (Glista and Scott, 1950; Wright, 1968; De Schrijver, 1977; Waldroup et al., 1985a).
Damage to the protein from overheating the soybean meal is more serious when dietary lysine concentrations are marginal,
and heat damage may be monitored by measuring the solubility of the protein, either by the Kjeldahl or by the dye-binding
method (Dale and Araba, 1987; Kratzer et al., 1990).
High level usage of soybean meal in poultry diets has been linked to the incidence of foot pad dermatitis (Jensen et al.,
1970). The exact cause of this is not known. Soybean meal contains relatively high levels of potassium, which may increase
litter moisture and thus result in sticky litter. In addition, the carbohydrate fraction of soybean meal is poorly digestible
(Parsons et al., 1980; Pierson et al., 1980) and may serve as a substrate for increased bacterial activity in the litter.
Animal Protein Sources
Animal protein sources—meat meals, fish meals, blood meal, and feather meal—are subject to variation as a result of
manufacturing conditions and the nature of the raw material from which they are processed. Excessive and/or prolonged
heating during drying will lower digestibility and cause some loss of essential amino acids. Proteins of hide, scales, hair,
feathers, and bone are not easily digested and contain high concentrations of keratin and/or collagenous proteins. The latter
will result in relatively low concentrations of tryptophan in the product. The use of certain lots of fish meal may result in the
development of a condition known as gizzard erosion (Janssen, 1971), a disease manifested primarily by ulcerations of the
lining of the gizzard. A substance known as gizzerosine has been isolated from samples of fish meal known to induce gizzard
erosion and has been shown to possess the same gizzard-erosion-producing properties (Okazaki et al., 1983). To date,
however, the exact level of gizzerosine necessary to induce gizzard erosion cannot be stated, since other factors (notably
excess levels of copper sulfate) may precipitate or exacerbate the condition.
Fish meal may result in the development of off-flavors in poultry meat (Fry et al., 1965) or eggs (Holdas and May, 1966;
Koehler and Bearse, 1975). The quantity of fish meal required to produce off-flavors is influenced primarily by the oil
content of the meal, length of time fed, degree of rancidity of the oil, and holding time and temperature of the egg or carcass.
Thus it is not possible to state a universal level of fish meal that will not result in the development of off-flavors.
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COMPOSITION OF FEEDSTUFFS USED IN POULTRY DIETS
71
ESTIMATING THE AMINO ACID COMPOSITION OF FEEDSTUFFS
Many factors influence the amino acid composition of grains and protein supplements. For accurate and economical feed
formulation, it is desirable to know the amino acid composition of the actual ingredient to be used in the diet. However, it is
generally not feasible to analyze all samples of feed ingredients prior to their use in feeds. Therefore research has been
conducted at several laboratories using regression analysis to estimate the amino acid composition of selected feed
ingredients from their proximate composition (Ward, 1989). An equation for estimating the amino acid content of feedstuffs
related to changes in protein content is presented in Table 9-4 and an equation for estimating amino acid content from other
proximate components is shown in Table 9-5. These equations represent different approaches that provide similar answers.
No attempts have been made to compare the results obtained from using both sets of equations on a common set of samples.
Knowledge of the availability of amino acids in feedstuffs is important for consistent formulation of diets that meet the
birds' amino acid requirements. The amounts of amino acids that are available to the animal are often much lower than the
quantity contained in feedstuffs. Many factors affect the availability of amino acids. Undenatured proteins vary markedly in
their digestibility. For example, feathers and most connective
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COMPOSITION OF FEEDSTUFFS USED IN POULTRY DIETS
TABLE 9-5 Estimation of Amino Acid Composition of Feed Ingredients from Proximate Components
Ingredients
Regression Factor
Methionine
Methionine +
Lysine
Threonine
Cystine
Lupin beans
Intercept
0.21996
0.95037
1.4019
0.25777
Protein
–a
–
0.018
0.02099
Moisture
-0.00306
-0.01326
-0.03354
-0.01034
Fat
0.0076
–
–
0.04113
Fiber
-0.00219
-0.01262
-0.0142
–
Ash
–
–
–
–
Milo
Intercept
0.0557
0.0859
0.2753
0.0593
Protein
0.0126
0.0282
0.0097
0.0238
Moisture
–
–
–
–
Fat
–
–
-0.0392
–
Fiber
–
0.0142
-0.0227
-0.014
Ash
–
-0.0237
0.0353
0.0318
Meat and bone
Intercept
0.7048
-1.1187
4.7627
-0.0022
meal
Protein
0.0098
0.0458
–
0.0384
Moisture
-0.0299
0.0372
-0.09
–
Fat
0.012
–
–
–
Fiber
0.0555
–
–
–
Ash
-0.0224
–
-0.0629
-0.0099
Poultry by-product
Intercept
-9.1947
8.587
-12.066
7.8878
Protein
0.1019
-0.0311
0.149
–
Moisture
0.1013
-0.0403
–
–
Fat
0.1438
-0.149
0.2488
-0.2065
Fiber
–
–
–
0.244
Ash
0.0801
-0.1338
0.1535
0.1618
Poultry byIntercept
0.9628
7.3812
11.8668
1.6665
product (crude
Protein
-0.0162
-0.0361
-0.0936
0.0137
protein = 54–62%)
Moisture
-0.0675
-0.1187
–
-0.042
Fat
0.0681
-0.1102
–
–
Fiber
0.0623
–
–
–
Ash
–
-0.0761
-0.1299
-0.0212
Field peas
Intercept
0.12772
0.18461
0.1614
0.39919
Protein
0.01941
0.04412
0.03032
-0.01403
Moisture
-0.00895
–
–
–
Fat
–
-0.05672
-0.11144
0.06006
Fiber
-0.01017
-0.01301
0.02799
0.01807
Ash
0.09637
–
0.12756
-0.10471
Rice bran (full-fat)
Intercept
0.0315
0.1517
-0.1305
0.0202
Protein
0.0135
0.0274
0.0313
0.0246
Moisture
–
–
–
0.0024
Fat
–
-0.0033
–
–
Fiber
–
-0.0046
–
0.0045
Ash
-0.0018
-0.0039
0.0061
0.001
Soybean meal
Intercept
0.1754
0.1902
-0.113
1.5584
(crude protein
Protein
0.0079
0.0179
0.0579
0.0159
=44–48%)
Moisture
–
–
–
-0.0289
Fat
–
–
–
-0.0366
Fiber
–
–
–
-0.0277
Ash
0.0221
0.0624
0.0665
–
Sunflower meal
Intercept
-0.0452
0.04425
1.1555
0.31712
Protein
0.01905
0.03874
0.0157
0.02928
Moisture
0.01612
0.00023
0.00358
Fat
–
–
–
-0.04026
Fiber
–
–
-0.01197
–
Ash
–
–
-0.03554
–
Wheat
Intercept
0.196
0.0074
0.3902
0.0717
Protein
0.0098
0.0582
0.0137
0.0336
Moisture
-0.0086
-0.0054
-0.0195
-0.0068
Fat
–
0.0435
0.0812
0.0545
Fiber
-0.0412
-0.0195
0.0163
0.0628
Ash
-0.0032
-0.0285
-0.0144
-0.0173
Intercept
0.0315
0.1517
-0.1305
0.0202
Bakery by-product
Protein
0.0315
0.0274
0.0313
0.0246
Moisture
–
–
–
0.0024
Fat
–
-0.0033
–
–
Fiber
–
-0.0046
0.0045
–
-0.0018
-0.0039
0.0061
0.001
Ash
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72
Tryptophan
Arginine
0.04185
0.010
–
–
–
–
0.142
0.014
0.0116
-0.07
–
-0.0637
-1.7233
0.0229
0.0562
0.0266
0.1311
–
0.8287
–
-0.0159
–
-0.055
-0.0079
0.0981
–
–
0.0257
–
0.0172
0.09402
0.12596
-0.02906
–
–
0.24338
0.0594
0.0042
–
–
–
0.0051
-0.201
0.0222
–
–
–
-0.0241
-0.35379
0.02035
0.7692
0.11352
-0.05846
–
–
-0.17185
0.2664
0.0163
0.0092
–
-0.0238
0.0741
5.4562
–
-0.0916
-0.0565
–
-0.0246
0.1536
0.0627
0.0423
–
–
–
2.4219
0.0306
–
0.06947
–
–
-0.21985
-0.0312
0.0433
–
–
–
–
1.0221
0.0678
–
–
–
-0.1132
-0.52833
0.09468
0.00528
0.0001
–
0.0582
0.0047
–
-0.0142
–
–
0.0594
0.0042
–
–
–
0.0051
–
–
–
0.381
0.0221
-0.0176
0.0154
–
-0.0016
-0.0312
0.0433
–
–
–
–
-0.0601
-0.91679
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COMPOSITION OF FEEDSTUFFS USED IN POULTRY DIETS
Ingredients
Regression Factor
Methionine
Barley
Intercept
Protein
Moisture
Fat
Fiber
Ash
Intercept
Protein
Moisture
Fat
Fiber
Ash
Intercept
Protein
Moisture
Fat
Fiber
Ash
Intercept
Protein
Moisture
Fat
Fiber
Ash
0.03751
0.01311
–
–
–
–
0.11324
0.01123
–
–
–
–
0.47972
0.02256
-0.01619
-0.00898
-0.05844
0.00788
8.8912
0.02597
–
–
-0.3727
-0.0272
Corn
Corn gluten meal
Fish meal
Methionine +
Cystine
-0.0319
0.02881
–
0.02886
0.01549
–
0.05313
0.02982
–
–
–
–
-0.05128
0.05079
-0.02883
-0.00663
–
0.00546
5.0029
–
-0.0651
-0.0702
–
-0.0754
73
Lysine
Threonine
Tryptophan
Arginine
0.05149
0.01975
0.01235
–
–
–
-0.10041
0.04573
–
–
–
–
-1.68796
0.04201
0.01719
-0.00561
0.12073
–
2.2017
0.055
0.06728
–
-0.7517
-0.0566
0.05491
0.02713
–
–
–
–
-0.05593
0.02275
0.00678
0.01593
0.00963
–
-1.42473
0.05376
–
0.00337
0.12052
-0.00359
4.4545
–
-0.0358
-0.03662
-0.182
-0.0612
0.00596
0.01053
–
–
–
–
0.26305
–
-0.01334
–
–
–
-3.55835
0.06078
–
-0.00604
0.22955
0.01117
-0.3998
0.0124
–
0.0241
-0.1369
0.009
-0.019
0.0339
0.01762
–
–
–
-0.03611
0.05484
–
–
–
–
-1.03918
0.04928
0.00518
-0.00384
0.04866
-0.0058
3.6336
0.02564
-0.0331
–
-0.2596
-0.0482
NOTE: To estimate amino acid, insert values shown for specific amino acid into the following equation: y = intercept + b1(% protein) + b2
(% moisture) + b3(% fat) + b4(% fiber) + b5(% ash), where the b, etc., represent the regression coefficients listed in each column. Dash
indicates that no coefficients were available.
Sources: This information is drawn from three reports published in 1986 by Monsanto: Amino Acids in Feed Ingredients and Their
Predictability. Monsanto Nutrition Update, vols. 4:2, 4:3, and 4:4. St. Louis, Mo.: Monsanto Company.
tissues contain high concentrations of cystine and disulfide bonding, which increase the stability of the protein and
resistance to digestive enzymes. Antinutritional factors such as tannins in sorghum and trypsin inhibitors in soybeans reduce
the availability of amino acids. Much of the latter adverse effect is due to increases in endogenous amino acid losses. The
negative effects of undenatured protein structure and antinutritional factors can usually be reduced or totally eliminated by
heat processing. Although some processing is needed to increase the availability of amino acids in many feedstuffs, adverse
processing conditions such as excessive pressure and heat can reduce availability. These factors are particularly critical for
animal protein meals since substantial processing or cooking is required during manufacturing. Lysine and cystine are two of
the amino acids most affected by processing conditions.
True digestibility coefficients for amino acids in 30 feedstuffs are shown in Table 9-6. The values were determined by
the precision-fed cockerel assay described by Sibbald (1986) or a modification thereof. The three primary sources of the
digestibility values used to compile the data of Table 9-6 were Sibbald (1986), Green (1987), and Parsons (1990a), with data
from other published reports also included. The assay was originally developed for determination of true ME (Sibbald, 1976)
and later extended to determination of amino acid digestibility (Likuski and Dorrell, 1978; Sibbald, 1979). The basic
procedure consists of subjecting adult male birds to fasting for 24 to 48 hours, followed by crop-intubation of 30 to 50 g of
the test feedstuff and quantitative collection of excreta for 48 hours. Additional cockerels are either subjected to fasting or
given a nitrogen-free diet during the assay period to estimate endogenous amino acid excretion. A large number of data have
been generated by using this assay during the last 10 years, and the results seem to be reasonably consistent among different
laboratories.
A large portion of the data used to derive the coefficients in Table 9-6 were determined with cecectomized birds;
however, data from studies with conventional birds were also included. Cecectomy removes the majority of the hindgut area
in poultry and eliminates most of the potentially confounding effects of the hindgut microflora on amino acid excretion. The
surgical procedure is simple, and several laboratories are currently using the technique. Digestibility coefficients determined
with cecectomized birds are often lower than those determined with conventional birds.
Determination of amino acid digestibility by analysis of the ideal contents has also been used to a limited extent. The
two primary approaches used in these studies
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COMPOSITION OF FEEDSTUFFS USED IN POULTRY DIETS
74
have been (1) removal of the ideal contents immediately following slaughter (Summers and Robblee, 1985) and (2) collection
of intestinal digesta via a cannula placed in the terminal ileum (Thomas and Crissey, 1983; Raharjo and Farrell, 1984).
It is generally accepted that digestible amino acid values are more indicative of relative nutritional value among
feedstuffs than are total amino acid concentration values. However, the application of digestibility values in practical feed
formulation is sometimes confusing because the amino acid requirements listed in the tables herein are expressed as total
amino acid concentration in the diet. There is little or no published research on the digestible amino acid requirements of
poultry species. Therefore a review of 28 published studies on the lysine and methionine plus cystine requirements of
broilers, turkeys, and laying hens was recently conducted to calculate digestible amino acid requirements indirectly (Parsons,
1990b). First, the amino acid digestibility coefficients in Table 9-6 were used to calculate the digestible amino acid content of
the basal diet feed ingredients used in the requirement studies. The digestible amino acid content of the basal diet was then
added to the amount of supplemental crystalline amino acid (100 percent available) needed to meet the requirement; this sum
was considered to be the digestible amino acid requirement. The results of these calculations for the 28 studies were
consistent and indicated that the calculated digestible amino acid requirements were 8 to 10 percent lower than the
determined total amino acid requirements.
Amino Acid Supplements
Individual amino acids are frequently included as ingredients in diets of poultry. DL-methionine and L-lysine are most
commonly used in commercial diets and other amino acids may be used in semipurified and purified diets. The protein
equivalents and estimated MEns of 20 amino acids are presented in Table 9-7. This information should be useful in
formulating poultry diets.
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TABLE 9-7 Nitrogen Concentration, Crude Protein Equivalents, and Nitrogen-Corrected Metabolizable Energy Values for Amino Acids
Amino Acid
Nitrogen (%)
Crude Protein Equivalent (g/100 g) of Amino Acid
Metabolizable Energy (kcal/kg)a
Alanine
15.72
98.25
3,060
Arginine
32.16
201.00
2,940
Asparagine
21.20
132.50
1,760
Aspartic acid
10.52
65.75
2,020
Cystine
11.66
72.88
2,060
Glutamic acid
9.52
59.50
2,880
Glutamine
19.17
119.81
2,630
Glycine
18.66
116.62
1,570
Histidine
27.08
169.25
2,410
Isoleucine
10.68
66.75
5,650
Leucine
10.67
66.69
5,640
Lysine
19.16
119.75
4,600
Methionine
9.39
58.69
3,680
Phenylalanine
8.48
53.00
6,030
Proline
12.17
76.06
3,980
Serine
13.33
83.31
2,210
Threonine
11.76
73.50
3,150
Tryptophan
13.72
85.75
5,460
Tyrosine
7.73
48.31
5,240
11.96
74.75
4,990
Valine
a
Assuming 100 percent digestibility and conversion of nitrogen to uric acid (including urea in the case of arginine).
CHARACTERISTICS OF DIETARY FATS
As discussed in Chapter 1, dietary fats vary appreciably in composition and in their contributions to nutrition of poultry.
The fatty acid composition of some ingredients commonly used in poultry diets is presented in Table 9-8. Selected
characteristics of supplemental fats (including combined moisture, insolubles, and unsaponifiables content), fatty acid
composition, and experimentally determined MEn values are shown in Table 9-9. This information provides an overview of
the different fats that have been evaluated experimentally and some of the conditions under which they were evaluated. For
comparative purposes, MEn values of specific carbohydrates are also listed in Table 9-9.
MACROMINERAL SUPPLEMENTS
Concentrated sources of calcium, phosphorus, sodium, potassium, and magnesium are often used to achieve desired
dietary concentrations of specific macrominerals. These mineral sources contain other elements of potential nutritional
importance, including chlorine, fluorine, sulfur,
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COMPOSITION OF FEEDSTUFFS USED IN POULTRY DIETS
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COMPOSITION OF FEEDSTUFFS USED IN POULTRY DIETS
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COMPOSITION OF FEEDSTUFFS USED IN POULTRY DIETS
78
iron, copper, manganese, and zinc. The concentration of these elements contained in selected macromineral supplements
is shown in Table 9-10.
MYCOTOXINS
Mycotoxins are toxic compounds produced by fungi. Most mycotoxins cause health problems for animals by entry
through the feed, although they may also be water- or air-borne. Given the appropriate conditions, fungi will grow on grain
and oilseeds prior to harvest. Wet conditions and warm temperatures favor the growth of fungi (Diener et el., 1987). Stresses
such as drought, insect infestation, and plant disease often make the crop susceptible to fungal growth. Some fungi will then
produce mycotoxins, which remain with the grain and oilseeds after harvest.
Mycotoxins in feed ingredients are difficult to economically remove or destroy. One method for detoxification of one
class of mycotoxins—aflatoxins—is ammoniation of ingredients. Ammoniation was effective in destroying aflatoxin in
peanut meal and cottonseed meal (Gardner et al., 1971) and in corn (Hughes et al., 1979). A second procedure for reducing
the effect of aflatoxins is the use of dietary adsorbents. Including sodium calcium aluminosilicate in the diet at a level of 0.5
percent is effective in reducing the effect of aflatoxins on the growth of chickens (Kubena et al., 1990).
Conditions that are favorable for fungal growth and mycotoxin production may also occur while ingredients are in
storage. The best way to prevent this problem is to keep the moisture level of ingredients low enough to inhibit fungal
growth. In some instances, antifungal additives may be used to prevent fungal growth in feed or ingredients.
Several classes of mycotoxins are known to cause economic losses in poultry. The first to be identified was aflatoxins.
These are produced by some strains of the fungi Aspergillus flavus, A. paraciticus , and A. nomius. Aflatoxins have been
labeled B1, B2, G1, and G2. Conditions appropriate for the production of aflatoxin are more commonly encountered in the
southeastern or central part of the United States or where the leaf canopy maintains high moisture content at the plant level.
Aflatoxins can produce a variety of effects. Broilers show decreased growth and increased kilogram feed:gain ratios
when fed 2.5 mg of aflatoxin per kilogram but not when fed 1.25 mg/kg (Smith and
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COMPOSITION OF FEEDSTUFFS USED IN POULTRY DIETS
79
Hamilton, 1970). When hens were fed diets with approximately 90 mg of aflatoxin per kilogram, egg production decreased
quickly and a high rate of mortality ensued (Hamilton, 1971). At a level of 1.5 mg/kg feed, aflatoxins caused fatty livers,
necrosis, and bile duct hyperplasia (Carnaghan et al., 1966). Hematological responses such as lowered serum protein, reduced
hemoglobin, and lower levels of serum triglycerides, phospholipids, and cholesterol result from moderate aflatoxin doses
(Tung et al., 1972).
Fusarium moniliforme is a fungus that may grow on grains. It is found to produce a thiaminase causing thiamin
deficiency in chicks (Fritz et al., 1973). Mortality is increased if additional thiamin is not supplied in contaminated diets.
Corn shown to contain F. moniliforme causes substantial mortality when fed to ducklings (Jeschke et al., 1987).
Tricothecenes constitute another group of fungal compounds that may decrease the performance of poultry. These
compounds may be produced by several genera of fungi but are most commonly metabolites of Fusarium . Laboratory
studies have shown that T-2 toxin at levels up to 20 mg/kg of diet may decrease weight gain and egg production (Wyatt et al.,
1973b, 1975). Oral lesions and digestive disturbances are caused by toxic concentrations of T-2.
Other tricothecenes produced by Fusarium are deoxynivalenol (DON), nivalenol, and diacetylnivalenol. These toxins
appear to be more toxic to swine, in which they may cause vomiting and feed refusal (Morehouse, 1985), than to poultry.
Adverse effects of Fusarium toxins on turkey reproduction have been reported (Allen et al., 1983).
Mycotoxins such as ochratoxin A and zearalenone have also been identified and may cause deleterious effects on
poultry. A review of their effects was done by the Council for Agricultural Science and Technology (1989).
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STANDARD REFERENCE DIETS FOR CHICKS
80
10
Standard Reference Diets for Chicks
Many laboratories that use Leghorn- or meat-type chicks for studies in animal behavior, biochemistry, microbiology,
nutrition, pathology, physiology, and toxicology need nutritionally complete standard reference diets. The use of standard
reference diets that are well defined facilitates more valid comparison of information obtained from experiments conducted
within and among laboratories. The diets shown in Table 10-1 have been used successfully in various laboratories and are
presented as guides to those requiring such formulations. The isolated soybean protein, casein, and chemically defined diets
contain some mineral and vitamin supplements not normally needed in practical diets.
Dextrose (glucose·H2O) rather than starch should be used in diets consisting primarily of purified intact proteins (such as
isolated soy protein and casein) to obtain improved performance. Diets containing substantial quantities of dextrose and
crystalline amino acids should be stored under refrigeration to minimize Maillard or Browning reactions. These chemically
defined diets are intended for short-term use (1 to 3 weeks) and will not support maximum growth over an extended period of
time.
Copyright © National Academy of Sciences. All rights reserved.
STANDARD REFERENCE DIETS FOR CHICKS
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81
STANDARD REFERENCE DIETS FOR CHICKS
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82
Appendixes
Copyright © National Academy of Sciences. All rights reserved.
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83
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84
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APPENDIXES
85
TABLE A–1 Documentation of Nutrient Requirements of Starting and Growing Leghorn—Type Chickens
Nutrient and Estimated
Age Period (Days)
Response Criteria
Breed
Requirement
Protein, %
20
0–14
Growth
White Leghorn
21.1
0–42
Growth
White Leghorn and
Rhode Island Red
14–20
84–140
Growth
White Leghorn
References
Grau and Kamei, 1950
Edwards et al., 1956
McNaughton et al.,
1977b
McNaughton et al.,
1977b
Leeson and Summers,
1979
Leeson and Summers,
1979
Leeson and Summers,
1979
Douglas and Harms, 1982
Maurice et al., 1982
15–18
0–42
Growth
White Leghorn
12
0–56
Growth
White Leghorn
16
56–84
Growth
White Leghorn
19
84–104
Growth
White Leghorn
14 and 21
12 or 13.6
56–140
0–42
Growth
Growth
16 or 13.6
42–140
Growth
18
18
12
16.5
22
0–28
0–42
42–140
140–504
0–28
Growth of muscle fiber
Growth
Growth
Laying
Growth
White Leghorn
Commercial brown-egg
layers
Commercial brown-egg
layers
White Leghorn
White Leghorn
White Leghorn
White Leghorn
White Leghorn
18
Isoleucine, %
0.5
Leucine, %
1.2
Lysine, %
0.9–1.1
0.94
0.70
<0.5
<0.45
0.68
Methionine, %
0.8
Methionine and cystine, %
0.8
0.59
0.45
Threonine, %
0.72
Valine, %
0.8
Requirements for
essential amino acids
described in review papers
Requirements for
essential amino acids
described in review papers
Requirements for
essential amino acids
described in review papers
Calcium
0.78
0–140
Growth
White Leghorn
Timson et al., 1983
Keshavarz, 1984
Keshavarz, 1984
Keshavarz, 1984
Leeson and Summers,
1984
Chi, 1985
8–18
Growth
White Leghorn
Mori and Okumura, 1984
8–18
Growth
White Leghorn
Mori and Okumura, 1984
0–42
1–21
35–49
56–98
98–147
0–504
Growth
Growth, feed efficiency
Growth, feed efficiency
Growth
Growth
Growth, egg production
White Leghorn
White Leghorn
White Leghorn
White Leghorn
White Leghorn
White Leghorn
Edwards et al., 1956
Chung et al., 1973
Chung et al., 1973
Berg, 1976
Berg, 1976
Keshavarz, 1984
0–14
Growth
White Leghorn
Grau and Kamei, 1950
0–14
0–504
0–42
Growth
Growth, laying
Growth
White Leghorn
White Leghorn
White Leghorn
Grau and Kamei, 1950
Keshavarz, 1984
Chi, 1985
7–21
Growth, feed efficiency
White Leghorn
Davis and Austic, 1982
8–18
Various
Growth
Growth
White Leghorn
Primarily White
Leghorn
Mori and Okumura, 1984
Almquist, 1952
Various
Growth
White Leghorn
Waldroup et al., 1980
Various
Growth, egg production
White Leghorn
Harms, 1984
0–153
Growth
White Leghorn
3.19
154–439
Egg production
White Leghorn
0.89
2.08
35–126
12–154
White Leghorn
White Leghorn
3.50
2.0–3.5
177–225
At 133 to 4th egg
White Leghorn
White Leghorn
Classen and Scott, 1982
Leeson et al., 1986
0.8
98–140
White Leghorn
Keshavarz, 1987
3.5
3.55
4.0
98–140
140–420
>112
Growth
Growth, subsequent egg
production
Egg production
Growth, bone
development
Growth, subsequent egg
production
Egg production
Egg production
Egg production
Hamilton and Cipera,
1981
Hamilton and Cipera,
1981
Classen and Scott, 1982
Classen and Scott, 1982
White Leghorn
White Leghorn
White Leghorn
Keshavarz, 1987
Keshavarz, 1987
Leeson and Summers,
1987b
Nonphytate phosphorus, %
0.4–0.6
0.25–0.30
0.31
7–28
0–140
112–140
Growth
Growth
Growth
White Leghorn
Brown-egg layers
White Leghorn
Gillis et al., 1949
Carew and Foss, 1980
Douglas and Harms, 1986
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Maurice et al., 1982
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APPENDIXES
86
Nutrient and Estimated
Requirement
Potassium, %
0.20–0.24
Age Period (Days)
Response Criteria
Breed
References
0–28
Growth, bone
calcification
White Leghorn
Gillis, 1948
Sodium, %
0.10–0.30
0.13
0.15
0–28
0–21
0–140
Growth
Growth
Growth
White Leghorn
White Rock
White Leghorn
Burns et al., 1953
Hurwitz et al., 1973
Manning and McGinnis,
1980
0–14
Growth, feed efficiency
Broiler Strain
Nam and McGinnis, 1981
0–140
Growth, sexual maturity
White Leghorn
Leeson and Summers,
1980
0–28
0–28
0–21
Deficiency, neuropathy
Growth
Growth
White Leghorn
Broiler strain
White Rock
Bird, 1949
Gardiner et al., 1960
Nugara and Edwards,
1963
0–140
0–28
Growth, perosis
Growth
New Hampshire
White Leghorn
Gallup and Norris, 1939a
Watson et al., 1971
0–42
White Rock
O'Dell et al., 1958
0–42
To 1st egg
0–7
7–21
Growth, feathering, bone
development
Growth
Growth, feed efficiency
Growth, feathering
Growth, feathering
White Rock
White Leghorn
White Leghorn
White Leghorn
Edwards et al., 1959
Rahman et al., 1961
Sunde, 1972
Sunde, 1972
0–56
0–56
0–21
0–28
Growth
Growth
Growth, feed efficiency
Growth
Rhode Island Red
Rhode Island Red
Broiler strain
New Hampshire
Hill and Matrone, 1961
Hill and Matrone, 1961
Waddell and Sell, 1964
Davis et al., 1968
Chlorine, %
0.13
Sodium chloride, %
0.25
Magnesium, mg/kg
300
250
594
Manganese, mg/kg
50
20
Zinc, mg/kg
35
20
20
78
52
Iron, mg/kg
40
4
56
75–80
Copper, mg/kg
4
Iodine, mg/kg
0.300
0–56
Growth
Rhode Island Red
Hill and Matrone, 1961
0–56
Growth, thyoid histology
Creek et al., 1957
0.400
0–56
Growth, thyoid histology
0.075
Selenium, mg/kg
0.01 to 0.05, depending
on dietary concentration
of Vitamin E
0.01 to 0.05, depending
on dietary concentration
of Vitamin E
Vitamin A, IU/kg
800–1600
0–35
Growth
White Leghorn and
Broiler strains
White Leghorn and
Broiler strains
Broiler strain
0–24
Growth
Plymouth Rock
Thompson and Scott,
1969
0–14
Growth
Plymouth Rock
Gries and Scott, 1972c
0–56
White Leghorn
Record et al., 1937
1,200–2,000
2,650
1,760–7,000
70–84
0–189
0–56
Growth, absence of
deficiency signs
Curative feeding
Growth
Growth
White Leghorn
White Leghorn
White Leghorn
4,400
0–113
Growth, E. acervulina
resistance
White Leghorn
Record et al., 1937
Taylor and Russell, 1947
Thornton and Whittet,
1962
Coles et al., 1970
Vitamin D3 IU/kg
180
0–84
Brown-egg layers
Baird and Greene, 1935
132
0–21
Broiler strain
McNaughton et al., 1977a
198
0–21
Broiler strain
McNaughton et al., 1977a
500
Adults
Growth, bone
development
Growth, bone
development
Growth, bone
development
Egg production, shell
quality
Various strains
Ameenuddin et al., 1985
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Creek et al., 1957
Rogler and Parker, 1978
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APPENDIXES
87
Nutrient and Estimated
Requirement
Vitamin E, IU/kg
60
Age Period (Days)
Response Criteria
Breed
References
Various
Various strains
Machlin and Gordon,
1962
30–50
Vitamin K, mg/kg
0.524–0.528
0.515
0–35
To prevent exudative
diathesis, encephalomalacia,
muscular degeneration
Growth
White Rock
Combs and Scott, 1974
0–28
0–84
Growth
Growth
White Rock
White Rock
0.524–0.528
0–28
Growth
White Rock
Nelson and Norris, 1960
Nelson and Norris,
1961a
Nelson and Norris,
1961b
Riboflavin, mg/kg
3.5 decreasing to 1.0
3.5 decreasing to 1.0
3
0–7
49–56
0–56
White Leghorn
White Leghorn
White Leghorn
2.3
0–42
Growth
Growth
Growth, prevention of curled
toe paralysis
Growth, prevention of curled
toe paralysis
Pantothenic acid, mg/kg
6
6.6
0–42
0–150
White Leghorn
New Hampshire
4.8
0–42
Growth
Growth, egg quality,
hatchability
Growth
0–56
42–77
0–28
Growth
Growth
Growth
Barred Plymouth Rock
White Leghorn
White Leghorn
Childs et al., 1952
Sunde, 1955
Patterson et al., 1956
0–77
0–23
0–42
0–21
Growth
Growth
Growth
Growth
White Leghorn
White Leghorn
White Leghorn
White Leghorn
Davis and Briggs, 1951
Ott, 1951
Miller et al., 1956
Patel and McGinnis,
1980
0–147
0–126
Growth, egg production
Growth
White Leghorn
White Leghorn
Nesheim et al., 1971
Tsiagbe et al., 1982
0–18
Growth, feed efficiency
Broiler strain
Anderson and
Warnick, 1970
0–35
0–28
0–35
Growth, feed efficiency
Growth
Growth
White Leghorn
Broiler strain
New Hampshire
March and Biely, 1955
Young et al., 1955
March and Biely, 1956
0–18
Growth
Broiler strain
Creek and Vasaitis,
1963
Thiamine, mg/kg
0.6–0.8
0–35
Growth
White Leghorn
0.88
0.88
0–28
0–28
Growth
Gain, feed efficiency
White Leghorn
White Leghorn
Arnold and Elvehjem,
1938
Thornton, 1960
Thornton and Shutze,
1960
Pyridoxine, mg/kg
2.8–3.0
5.7
5
0–28
0–56
0–21
Growth
Growth
Growth
White Leghorn
White Plymouth Rock
Broiler strain
Niacin, mg/kg
28
1.8
17.5–20
Vitamin B12, mg/kg
4.4
27
2.5
10
Choline, mg/kg
2,000
1,000
Biotin, µg/kg
260
Folic Acid, mg/kg
0.80
0.30
0.33 to 1.45, depending
on protein level
0.30
White Leghorn
White Leghorn
Copyright © National Academy of Sciences. All rights reserved.
Heuser et al., 1938
Heuser et al., 1938
Bethke and Record,
1942
Bootwalla and Harms,
1990
Bauernfeind et al., 1942
Balloun and Phillips,
1957b
Bootwalla and Harms,
1991
Briggs et al., 1942
Fuller and Kifer, 1959
Kazemi and Kratzer,
1980
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APPENDIXES
88
TABLE A–2 Documentation of Nutrient Requirements of Leghorn—Type Chickens in Egg Production
Nutrient and Estimated
Age Period (Weeks)
Response Criteria
Breed
Requirement
Protein, g/bird daily
14.9
24–60
Egg yield
White Leghorn
14
15
Arginine, mg/bird daily
400
Isoleucine, mg/bird daily
475
650
Lysine, mg/bird daily
690
References
24–72
20–72
Egg yield
Egg yield
White Leghorn
White Leghorn
Balloun and Speers,
1969
Thayer et al., 1974
Proudfoot et al., 1988
Not specified
Egg yield
White Leghorn
Adkins et al., 1962
Not specified
Not specified
Egg yield
Egg yield
White Leghorn
White Leghorn
Bray, 1969
Gous et al., 1987
22–42
Egg yield
White Leghorn
Egg yield
Egg yield
White Leghorn
White Leghorn
Nathanael and Sell,
1980
Latshaw, 1981
Proudfoot et al., 1988
Egg yield
Egg yield
White Leghorn
White Leghorn
Reid and Weber, 1973
Latshaw, 1981
Egg yield
White Leghorn
Adkins et al., 1958
Egg yield
Egg yield
White Leghorn
Rhode Island Red
Wethli and Morris, 1978
Ohtani et al., 1989
Egg yield
Crossbreds
Hurwitz and Bornstein,
1978
650
24–72
620
20–72
Methionine + cystine, mg/bird daily
500
20 from onset of lay
530
24–72
Threonine, mg/bird daily
400
Not specified
Tryptophan, mg/bird daily
165
20–76
239
20–76
Valine, %
0.64
Not specified
Linoleic acid, %
2.0
1.0
1.0
0.9
Calcium, g/bird daily
3.12
22–54
22–54
22–54
20–72
Egg production
Egg weight
Hatch
Egg weight
White Leghorn
White Leghorn
White Leghorn
White Leghorn
Menge, 1970
Menge, 1970
Menge, 1970
Whitehead, 1981
48–55
White Leghorn
Atteh and Leeson, 1983
3.15
24–72
Egg production, shell
strength
Egg production
White Leghorn
>2.8
54–58
Egg production, shell
strength
White Leghorn
Scheideler and Sell,
1986
Austic and Keshavarz,
1988
Egg production
Egg production
Egg production
Egg production
White Leghorn
White Leghorn
White Leghorn
White Leghorn
Miles et al., 1983
Said and Sullivan, 1985
Sell et al., 1987
Sell et al., 1987
Egg production, egg
weight, shell thickness
White Leghorn
Leach, 1974
White Leghorn
Reid, 1977
Medium weight brownegg layers
Sauveur and Mongin,
1978
Nonphytate Phosphorus, mg/bird daily
215
28–36
250
21–32
250
35–51
>150
52–72
Potassium, %
0.10
12
Sodium, mg/bird daily
140–150
20–48
130
21–45
Egg production, feed
conversion
Egg yield
Chlorine, mg/bird daily
132
Magnesium, mg/kg
350
Not specified
Egg production
White Leghorn
Vogt, 1977
25–31
White Leghorn
Cox and Sell, 1967
900
Not specified
Egg production, egg
weight
Egg production
White Leghorn
355
30–38
Egg production,
hatchability
White Leghorn
Edwards and Nugara,
1968
Hajj and Sell, 1969
Managenese, mg/kg
>13
21–33
New Hampshire
20
22
>7
Zinc, mg/kg
28
54
17–23
Egg production,
hatchability
Egg production, egg
weight, shell quality
Shell quality
Iron, mg/kg
45
55
White Leghorn
Gallup and Norris,
1939b
Cox and Balloun, 1969
White Leghorn
Longstaff and Hill, 1971
22–72
Not specified
Egg yield, hatchability
Feather condition of
progeny
White Leghorn
White Leghorn
Stahl et al., 1986
Stahl et al., 1986
Not specified
Not specified
Hematocrit
Hatchability
White Leghorn
White Leghorn
Morck and Austic, 1981
Morck and Austic, 1981
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APPENDIXES
Nutrient and Estimated
Requirement
Copper, mg/kg
>1
<2.5
Iodine, µg/kg
35
>75
Selenium, mg/kg
0.05
0.05
89
Age Period (Weeks)
Response Criteria
Breed
References
44–48
44–48
Shell quality
Shell quality
White Leghorn
White Leghorn
Baumgartner et al., 1978
Baumgartner et al., 1978
4–45
4–45
Hatchability
Embryonic thyroid
White Leghorn
White Leghorn
Rogler et al., 1959a
Rogler et al., 1959b
32–56
32–57
Egg production
Egg production,
hatchability
White Leghorn
White Leghorn
Latshaw et al., 1977
Combs and Scott, 1979
Vitamin A, IU/kg
3,520
26–70
White Leghorn
Hill et al., 1961
2,750
20–64
Egg production, blood
spots, hatchability
Egg production, fertility,
hatchability
White Leghorn
Reid et al., 1965
Vitamin D3, IU/kg
150
21–34
White Leghorn
Abdurahim et al., 1979
250
30–46
Egg production, shell
quality, fertility,
hatchability
Egg production, shell
quality
White Leghorn
Shen et al., 1981
Not specified
Not specified
Hatchability
Hatchability
White Leghorn
White Leghorn
Jensen and McGinnis, 1960
Olson et al., 1962
Not specified
Hatchability
White Leghorn
Griminger, 1964
30–45
30–45
Egg production
Hatchability, chick quality
White Leghorn
White Leghorn
Petersen et al., 1947a
Petersen et al., 1947b
Not specified
Not specified
28–53
28–53
28–53
Hatchability
Hatchability
Egg production
Hatchability
Viability of progeny
White Leghorn
New Hampshire
White Leghorn
White Leghorn
White Leghorn
Gillis et al., 1948
Balloun and Phillips, 1957a
Beer et al., 1963
Beer et al., 1963
Beer et al., 1963
Not specified
White Leghorn
Ringrose et al., 1965
White Leghorn
Ringrose et al., 1965
White Leghorn
Ouart et al., 1987
Vitamin E, IU/kg
12
41 in presence of oxidized
fat
Vitamin K, mg/kg
>1.0
Riboflavin, mg/kg
2.5
3.6
Pantothenic acid, mg/kg
6.5
7
1.9
4.9
8.9
Niacin, mg/kg
9
11
Not specified
<21
Vitamin B12, µg/kg
1.0
41–57
Egg production,
hatchability
Egg production,
hatchability
Egg yield, hatchability
22–35
Hatchability
White Leghorn
Not specified
Not specified
Hatchability
Hatchability
New Hampshire
White Leghorn
Mariakulandai and
McGinnis, 1953
Johnson, 1954
Chin et al., 1958
50–66
45–57
32–52
Egg yield
Egg yield
Egg yield
White Leghorn
White Leghorn
White Leghorn
Miles et al., 1986
Parsons and Leeper, 1984
Keshavarz and Austic, 1985
19–73
Egg production
White Leghorn
Whitehead, 1980
44–55
White Leghorn
Sunde et al., 1950a,b
Not specified
Egg production,
hatchability
Hatchability
White Leghorn
Couch and German, 1950
1–2
0.5–1.0
Choline, mg/kg
1,050
<1,480
1,000
Biotin, mg/kg
0.10
Folic acid, mg/kg
0.5
0.2
Thiamin, mg/kg
0.68
Pyridoxine, mg/kg
2.5
Not specified
Hatchability
White Leghorn
Polin et al., 1963
Not specified
White Leghorn
Cravens et al., 1946
2.3
Not specified
White Leghorn
Fuller et al., 1961
4.5
Not specified
Egg production,
hatchability
Egg production,
hatchability
Egg production,
hatchability
White Leghorn
Fuller et al., 1961
Copyright © National Academy of Sciences. All rights reserved.
APPENDIXES
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APPENDIXES
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APPENDIXES
Nutrient and Estimated
Requirement
0.3–0.45 mg
0.34–0.49 mg
Thiamin, mg/kg
0.75
96
Age Period (Days)
Response Criteria
Breed
References
1–20
1–28
Growth, perosis
Growth, leg abnormalities
Arbor Acres
Not specified
Creek and Vasaitis, 1963
Saxena et al., 1954
3–28
Growth, polyneuritis
Thornton 1960
1.0–1.3
Not specified
Growth, feed efficiency
New Hampshire ×
Delaware
New Hampshire ×
Delaware
Thornton and Shutze
1960
Pyridoxine, mg/kg
3–5
12–42
White Rock
Hogan et al. 1941
2
<5.7
3.3
7–28
1–56
1–14
Growth, perosis, anemia,
dermatitis
Growth, feed efficiency
Growth, feed efficiency
Growth
Not specified
White Plymouth Rock
White Plymouth Rock
2.2–2.6
1–28
Kratzer et al., 1947
Fuller and Kifer, 1959
Fuller and Dunahoo,
1959
Daghir and Balloun, 1963
2.8–3.6
Vantress × Arbor Acre
1–14 or 35
Growth, gizzard erosion,
serum glutamic oxaloacetic
transaminase
Growth, feed efficiency
3
Not specified
Growth, feed efficiency
Not specified
>3.1
7–28
Not specified
3.2–3.4
1–28
Growth, serum aspartate
aminotransferase
Growth, perosis
≤1.0
1.1
1–20
8–17
Growth, feed efficiency
Growth
1.75
3–49
Growth, plasma amino acids
White Plymouth
Rock × Vantress
Ross
New Hampshire ×
Columbian
Not specified
1.3–2.7
1–21
Growth
Not specified
≤1.48
1–49
Growth
Not specified
Not specified
Copyright © National Academy of Sciences. All rights reserved.
Kirchgessner and
Friesecke, 1963
Maier and Kirchgessner,
1968
Daghir and Shah, 1973
Gries and Scott, 1972a
Lee et al., 1976
Yen et al., 1976
aboaysha and Kratzer,
1979
Kazemi and Kratzer,
1980
Blalock et al., 1984
APPENDIXES
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APPENDIXES
98
Nutrient and Estimated
Requirement
Sodium, mg/bird daily
<154
Age Period (Weeks)
Response Criteria
Breed
References
32–64
Egg production egg weight,
fertility, egg specific gravity,
hatchability
Cobb
Damron et al., 1983
Chlorine, mg/bird daily
208
32–60
Egg production, egg weight,
hatchability
Cobb
Harms and Wilson, 1984
Biotin, µg/bird daily
16
20–58
Egg production, egg weight,
hatchability
Marshall
Whitehead et al., 1985
TABLE A-5 Documentation of Nutrient Requirements of Broiler Breeder Males
Age Period (Days)
Response Criteria
Nutrient and Estimated
Requirement
Metabolizable energy, kcal/bird daily
400
28–40
Body weight, fertility,
hatchability, chick
production, testes weight
458
30–54
Body weight, fertility,
hatchability, chick
production, testes weight
346
30–46
Body weight, fertility,
hatchability, chick
production, testes weight
358
30–60
Body weight, semen
volume, sperm cells,
fertility
Protein, %
12.4
7–21
Development of testes,
subsequent fertility
12–14
4–53
Weight gain, semen
volume and concentration
testes weight
9
6–53
Weight gain, semen
volume and concentration
testes weight
15
1–4
Fertility 24–27 weeks
Protein, g/bird daily
10–14
20–60
Semen production
Breed
References
Broiler strain
McCartney and Brown,
1980
Broiler strain
Brown and McCartney,
1983
Hubbard
Brown and McCartney,
1986
Broiler strain
Buckner et al., 1986
Peterson
Wilson et al., 1971
Broiler strain
Wilson et al., 1987a
Broiler strain
Wilson et al., 1987b
Hubbard
Vaughters et al., 1987
Hubbard
Buckner and Savage,
1986
Calcium, %
<0.2
36–60
Semen volume, sperm
concentration, dead sperm,
fertility, hatchability
White Leghorn
Wilson et al., 1969
Calcium, mg/bird daily
7.98
44–56
Weight gain, blood
parameters, bone
constituents
Reproductive parameters
White Leghorn
Norris et al., 1972
Broiler strains
Kappleman et al., 1982
Weight gain, blood
parameters, bone
constituents
White Leghorn
Norris et al., 1972
Semen volume
Arbor Acres, cage
males
Bootwalla and Harms,
1989
<500
Nonphytate phosphorus, %
0.1
Not specified
44–56
Nonphytate phosphorus, mg/bird daily
32–40
110
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APPENDIXES
99
TABLE A-6 Documentation of Nutrient Requirements of Turkeys
Nutrient and Estimated
Age Period (Days)
Response Criteria
Requirement
Protein, %
28
0–7
Growth
20
0–4
Growth
20
8–16
Growth
28
0–8
Growth
25–32
0–6
Growth
18
8–12
Growth
16
12–16
Growth
14
16–20
Growth
22
8–12
Growth
18
12–16
Growth
14
16–20
Growth
24
8–10
Growth
20
10–12
Growth
18
12–14
Growth
24
6–12
Growth
30
0–7
Growth
22
7–13
Growth
30
0–4
Growth
21.3
10
Growth
19.5
14
Growth
17.6
18
Growth
21.7
10
Growth
18.4
14
Growth
15.0
18
Growth
20
5–14
Growth, carcass
composition
26
4–10
Growth, carcass quality
20
10–13
Growth, carcass quality
18
10–13
Growth, carcass quality
Arginine, %
1.60
0–3
Growth
1.90
1–3
Growth
1.60
1–3
Growth
Breed
References
Bronze, both sexes
Jersey Buff, both sexes
Large White, both sexes
Bronze, both sexes
Bronze, both sexes
Bronze, females
Bronze, females
Bronze, females
Large White, males
Large White, males
Large White, males
Large White, females
Large White, females
Large White, females
Large White, males
Large White, males
Large White, males
Large White, males
Large White, males
Large White, males
Large White, males
Large White, females
Large White, females
Large White, females
Large White, both sexes
Lloyd et al., 1949
Baldini et al., 1954
Carter et al., 1957
Atkinson et al., 1957
Balloun et al., 1959
Jensen et al., 1965
Jensen et al., 1965
Jensen et al., 1965
Summers et al., 1968
Summers et al., 1968
Summers et al., 1968
Summers et al., 1968
Summers et al., 1968
Summers et al., 1968
Eberst et al., 1972
Herz et al., 1975a
Herz et al., 1975b
Richter et al., 1980
Potter et al., 1981
Potter et al., 1981
Potter et al., 1981
Potter et al., 1981
Potter et al., 1981
Potter et al., 1981
Richter and Prinz, 1980
Small White, both sexes
Small White, males
Small White, females
Salmon, 1984
Salmon, 1984
Salmon, 1984
Almquist, 1952
Dunkelgod et al., 1970
Warnick and
Anderson, 1973
D'Mello and Emmans,
1975
Hurwitz et al., 1983a
1.75
1–3
Growth
Bronze, both sexes
Bronze, both sexes
Bronze and Large White,
both sexes
Large White, males
1.59
0–4
1.32
4–8
1.02
8–12
0.80
12–16
0.63
16–20
0.47
20–24
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Glycine, %
0.90
0–3
Growth
Bronze, both sexes
Kratzer and Williams,
1948a
Histidine, %
0.58
1–3
Growth
Bronze, both sexes
0.53
0–4
0.42
4–8
0.30
8–12
0.23
12–16
0.18
16–20
0.12
20–24
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males
mathematical model
Large White, males,
mathematical model
Warnick and
Anderson, 1973
Hurwitz et al., 1983
Copyright © National Academy of Sciences. All rights reserved.
Hurwitz et al., 1983a
Hurwitz et al., 1983a
Hurwitz et al., 1983a
Hurwitz et al., 1983a
Hurwitz et al., 1983a
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
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APPENDIXES
100
Nutrient and Estimated
Requirement
Isoleucine, %
0.80
1.10
Age Period (Days)
Response Criteria
Breed
References
0–3
1–3
Growth
Growth
Bronze, both sexes
Bronze, both sexes
0.84
1.03
1–3
0–4
0.86
4–8
0.67
8–12
0.53
12–16
0.42
16–20
0.31
20–24
Growth
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Large White, males
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Kratzer et al., 1952
Warnick and
Anderson, 1973
D'Mello, 1975
Hurwitz et al., 1983
Leucine, %
1.86
1–3
Growth
Bronze, both sexes
1.42
1.96
1–3
0–4
1.62
4–8
1.23
8–12
0.96
12–16
0.74
16–20
0.53
20–24
Growth
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Large White, males
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Lysine, %
1.5
0.96
0.85
0.76
0.56
0.60
1.55
0–4
4–8
8–12
14–18
16–19
20–23
0–6
Growth
Growth
Growth
Growth
Growth
Growth
Growth
Bronze, both sexes
Bronze, both sexes
Bronze, both sexes
Bronze, both sexes
Bronze, both sexes
Bronze, both sexes
Bronze, both sexes
1.60
1.68
0–3
1–3
Growth
Growth
Large White, both sexes
Bronze, both sexes
1.50
0–4
Growth
Large White, males
1.40
4–8
Growth
Large White, males
1.12
8–12
Growth
Large White, males
1.55
1–3
Growth
Large White, males
0.96
0.76
1.4
1.2
0.9
1.42
12–16
16–20
8–12
12–16
11–20
0–4
1.12
4–8
0.81
8–12
0.63
12–16
0.49
16–20
0.32
20–24
Growth
Growth
Growth
Growth
Growth
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Large White, males
Large White, males
Large White, both sexes
Large White, both sexes
Large White, both sexes
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Methionine, %
0.55
0.56
Starting
0–6
Growth
Growth
Bronze, both sexes
Jersey Buff, both sexes
Copyright © National Academy of Sciences. All rights reserved.
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Warnick and
Anderson, 1973
D'Mello, 1975
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Almquist, 1952
Kratzer et al., 1956b
Kratzer et al., 1956b
Kratzer et al., 1956b
Kratzer et al., 1956b
Kratzer et al., 1956b
Balloun and Phillips,
1957b
Kummero et al., 1971
Warnick and
Anderson, 1973
Tuttle and Balloun,
1974
Tuttle and Balloun,
1974
Tuttle and Balloun,
1974
D'Mello and Emmans,
1975
Jensen et al., 1976
Jensen et al., 1976
Potter et al., 1981
Potter et al., 1981
Potter et al., 1981
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Almquist, 1952
Baldini et al., 1957
APPENDIXES
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APPENDIXES
102
Nutrient and Estimated
Requirement
0.88
Age Period (Days)
Response Criteria
Breed
References
4–8
8 – 12
0.53
12 – 16
0.41
16 – 20
0.30
20 – 24
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Hurwitz et al., 1983
0.67
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Threonine, %
1.10
1.00
1–2
1–3
Growth
Growth
Large White, males
Bronze, both sexes
0.94
1.14
1–3
0–4
0.94
4–8
0.72
8 – 12
0.56
12 – 16
0.44
16 – 20
0.32
20 – 24
Growth
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Large White, males
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Tryptophan, %
0.26
0.37
0.26
0–4
1–2
1–3
Growth
Growth
Growth
Bronze, both sexes
Large White, males
Bronze, both sexes
0.21
0–4
0.17
4–8
0.13
8 – 12
0.11
12 – 16
0.08
16 – 20
0.06
20 – 24
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Valine, %
1.38
1.20
1–2
1–3
Growth
Growth
Large White, males
Bronze, both sexes
1.21
1.34
1–3
0–4
1.13
4–8
0.88
8 – 12
0.69
12 – 16
0.53
16 – 20
0.40
20 – 24
Growth
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Carcass content plus
maintenance
Large White, males
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Large White, males,
mathematical model
Linoleic, %
1.00
0–3
Growth
Large White and Bronze,
both sexes
Ketola et al., 1973
Calcium, %
1.7
0–3
Bone ash
Bronze, both sexes
1.5
0–4
Bone ash
Small White, both sexes
Motzok and Slinger,
1948
Wilcox et al., 1953
Copyright © National Academy of Sciences. All rights reserved.
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Dunkelgod et al., 1970
Warnick and
Anderson, 1973
D'Mello, 1976
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Almquist, 1952
Dunkelgod et al., 1970
Warnick and
Anderson, 1973
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Dunkelgod et al., 1970
Warnick and
Anderson, 1973
D'Mello, 1975
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
Hurwitz et al., 1983
APPENDIXES
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103
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APPENDIXES
104
Nutrient and Estimated
Requirement
Riboflavin, mg/kg
2.7
Age Period (Days)
Response Criteria
Breed
References
0–6
Bronze, both sexes
Patrick et al., 1944
3.75
0–4
Bronze, both sexes
Bird et al., 1946
4.0
0–6
0–3
>3.50
0–3
Bronze and Large
White, both sexes
Medium White, both
sexes
Large White, both
sexes
Jukes et al., 1947
4.0
Growth, deficiency
symptoms
Growth, deficiency
symptoms
Growth, deficiency
symptoms
Erythrocyte glutathione
reductase and liver flavin
Growth, leg paralysis
Pantothenic acid, mg/kg
10.5
1–3
Growth, dermatitis
Bronze, both sexes
<8.6
0–3
Growth
Large White, both
sexes
Kratzer and Williams,
1948b
Ruiz and Harms, 1989b
Niacin, mg/kg
71.5
21
0–2
4–12
Growth, enlarged hocks
Growth, leg disorders
Scott, 1953
Christmas et al., 1986
44
0–3
Growth, leg disorders
Bronze, both sexes
Large White, both
sexes
Large White, both
sexes
Vitamin B12, mg/kg
0.002–0.010
0–4
Growth
Bronze, both sexes
0.003
0–6
Growth
Small White, both
sexes
Sherwood and Sloan,
1954
Johnson, 1955
Choline, mg/kg
2,000
1,900
2,300
<1,490
0–2
0–6
10–24
0–3
Perosis
Perosis
Growth
Growth
Jukes, 1940
Evans, 1943
Slinger et al., 1946
Harms and Miles, 1984
<1,250
4–8
Growth
Not specified
Not specified
Bronze, females
Large White, both
sexes
Large and Medium
White, both sexes
Biotin, mg/kg
0.284
0–3
Bronze, both sexes
0.275–0.324
0–3
Bronze, both sexes
Jensen and Martinson,
1969
Dobson, 1970
0.225–0.275
0–3
Bronze, both sexes
Dobson, 1970
0.220
Folic acid, mg/kg
0.8
0–8
Growth, deficiency
symptoms
Growth, deficiency
symptoms
Growth, deficiency
symptoms
Growth
Large White, males
Krueger et al., 1976
Growth, anemia
prevention
Growth, cervical paralysis
Bronze, both sexes
Jukes et al., 1947
Jersey Buff, both sexes
Russell et al., 1947
Bronze, both sexes
Robenalt, 1960
0–3
Growth, symptoms of
deficiency
Growth
Bronze, both sexes
Sullivan et al., 1967
0–3
0–4
Growth
Growth, survival
Not specified
Bronze, both sexes
Kratzer et al., 1947
Sullivan et al., 1967
2.0
Thiamin, mg/kg
2.0
1.6–2.0
Pyridoxine, mg/kg
2.0–3.0
3.9–4.4
0–6
0–3
0–3
Copyright © National Academy of Sciences. All rights reserved.
Lee, 1982
Ruiz and Harms, 1989a
Ruiz and Harms, 1989b
Blair et al., 1986
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APPENDIXES
105
TABLE A-7 Documentation of Nutrient Requirements of Turkey Breeders
Nutrient and Estimated
Age Period (Days)
Response Criteria
Requirement
Protein, %
15
32–52
Poult yield
15
30–48
Poult yield
10
18
14
14
12
14
10
8
16
Protein, g/bird daily
26
Linoleic acid, %
1.21
30–46
32–48
17–20
20–32
12–28
28–56
30–41
28–53
32–48
1.1
Calcium, %
1.75
2.0
Breed
References
Large White, females
Jensen and McGinnis,
1961
Atkinson et al., 1970
Poult yield
Poult yield egg weight
Egg production
Egg production
Semen production
Egg production
Poult yield
Semen production
Poult yield
Large and Small
White, females
Large White, females
Large White, females
Large White, females
Large White, females
Large White, males
Large White, females
Large White, females
Large White, males
Large White, females
Minear et al., 1972
Menge et al., 1979
Meyer et al., 1980a
Meyer et al., 1980a
Meyer et al., 1980b
Meyer et al., 1980a
Meyer et al., 1980b
Cecil, 1982
Bougon et al., 1985
32–60
Poult yield
Small White, females
Jackson et al., 1974
24–55
Large White, females
30–55
Egg production,
hatchability
Poult yield
Large White, females
Cooper and Barnett,
1968
Whitehead and Herron,
1988
26–54
30–48
Poult yield
Poult yield
Bronze, females
Large White, females
30–47
30–47
30–47
30–46
0–4
33–53
30–50
Egg production
Egg production
Egg production
Poult yield
Growth
Poult yield
Poult yield
Bronze, females
Large White, females
Bronze, females
Large White, females
Large White, males
Small White, males
Large White, females
30–42
30–50
30–45
30–50
Poult yield
Poult yield
Poult yield
Fertility
Small White, females
Large White, females
Small White, females
Medium White,
females
Ferguson et al., 1974
Waldroup et al., 1974
Atkinson et al., 1976
Slaugh et al., 1989
30–46
Poult yield
Bronze, females
Atkinson et al., 1967b
30–48
Hatchability, poult survival
Bronze, females
Jensen, 1965
32–40
31–40
Poult yield
Poult yield
Bronze, females
Large White, females
300–400
41–53
Poult yield
Large White, females
900
29–35
Adequate poult yield but
inadequate liver storage
Large White, females
Wilhelm et al., 1941
Kramer and Waibel,
1978
Kramer and Waibel,
1978
Stevens et al., 1984
Vitamin E, IU/kg
24
32–54
Poult yield
Bronze, females
Jensen and McGinnis,
1957
Not specified
Poult yield
Bronze, females
Boucher et al., 1942
Various
Poult yield, survival
Bronze, females
Kratzer et al., 1955
32–48
Egg weight, poult yield
Large White, females
Harms et al., 1988
32–46
Poult yield
<1,230
Biotin, mg/kg
>0.105
<0.150
32–54
Poult yield
Bronze and Large
White, females
Small White, females
Balloun and Miller,
1964a
Ferguson et al., 1975
30–46
Not specified
Poult yield
Poult yield
Waibel et al., 1969
Arends et al., 1971
0.160
27–34
Egg biotin (albumen)
Large White, females
Large and Medium
White, females
Medium White,
females
Folic acid, mg/kg
0.7
1.23
32–48
32–48
Poult yield
Poult yield, survival
1.9
2.66
3.19
2.25
1.2
2.5
2.55
Nonphytate phosphorus, %
0.42
0.30
0.55
0.30
Manganese, mg/kg
60
Vitamin A, IU/kg
2,200–3,520
Vitamin D, IU/kg
1,000
<750
Riboflavin, mg/kg
3.50
Pantothenic acid, mg/kg
16.0
Niacin, mg/kg
23.6
Choline, mg/kg
<990
Bronze, females
Large White, females
Copyright © National Academy of Sciences. All rights reserved.
Jensen et al., 1963
Balloun and Miller,
1964b
Atkinson et al., 1967a
Atkinson et al., 1967a
Atkinson et al., 1967a
Arends et al., 1967
Neagle et al., 1968
Potter et al., 1974
Waldroup et al., 1974b
White et al., 1987
Kratzer et al., 1956a
Miller and Balloun, 1967
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APPENDIXES
106
TABLE A-8 Documentation of Nutrient Requirements of Geese
Nutrient and Estimated
Age Period (Days)
Response Criteria
Requirement
Protein, %
24
0–6
Growth
Breed
References
White Chinese
12
6–16
Growth
White Chinese
24
0–4
Growth
White Chinese
16
4–12
Growth
White Chinese
20
16
14
18.2
12.0
18
0–4
4–6
4–9
0–2
2–7
0–3
Embden
Embden
Embden
Not specified
Not specified
Embden
16
0–9
Growth, feathering
Growth, feathering
Growth, feathering
Growth, feed efficiency
Growth, feed efficiency
Growth, carcass yield,
carcass composition
Growth, carcass yield,
carcass composition
Roberson and Francis,
1963a
Roberson and Francis,
1963a
Roberson and Francis,
1963b
Roberson and Francis,
1963b
Allen, 1981
Allen, 1981
Allen, 1981
Nitsan et al., 1983
Nitsan et al., 1983
Summers et al., 1987
Embden
Summers et al., 1987
1–2 and 3–7
0–4
4–8
0–2
2–7
Growth
Growth
Growth
Growth, feed efficiency
Growth, feed efficiency
White Chinese
Not specified
Not specified
Not specified
Not specified
Roberson and Francis, 1966
Mateova et al., 1980
Mateova et al., 1980
Nitsan et al., 1983
Nistan et al., 1983
0–3
Growth, feed efficiency,
carcass composition
Growth, feed efficiency
Growth, feed efficiency
White Italian
Znaniecke et al., 1975
Not specified
Not specified
Nitsan et al., 1983
Nitsan et al., 1983
White Italian
Znaniecka et al., 1975
0–2
2–7
Growth, feed efficiency,
carcass composition
Growth, feed efficiency
Growth, feed efficiency
Not specified
Not specified
Nitsan et al., 1983
Nitsan et al., 1983
0–4 and 0–6
Growth, bone ash
Pilgrim
Aitken et al., 1958
0–4 and 0–6
Growth, bone ash
Pilgrim
Aitken et al., 1958
0–2
Growth
Embden
Serafin, 1981
0–3
Growth, mortality
Embden
Serafin, 1981
0–3
0–3
Growth, perosis
Growth
Not specified
Embden
Battig et al., 1953
Serafin, 1981
0–3
Growth, perosis
Embden
Serafin, 1981
0–2
Growth, liveability
Toulouse
Briggs et al., 1953
Lysine, %
0.90
1.10
0.85
1.07
0.60
Methionine, %
0.40
0.29
0.15
Methionine + cystine, %
0.73
0.58
0.47
Calcium, %
0.4
Total phosphorus, %
0.46
Riboflavin, mg/kg
3.8
Pantothenic acid, mg/kg
12.6
Niacin, mg/kg
66
31.2
Choline, mg/kg
1530
Choline, niacin, folic acid
Not determined but
estimates obtained
0–2
2–7
0–3
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APPENDIXES
107
TABLE A-9 Documentation of Nutrient Requirements of Ducks
Nutrient and Estimated
Age Period (Days)
Response Criteria
Requirement
Protein, %
22
0–2
Growth
16
2–7
Growth
18
0–2
Growth
16
2 to market
Growth
19
0–2
Growth
16
3–8
Growth
Arginine, %
1.08
1–3
Growth, feed efficiency
Isoleucine, %
0.63
1–3
Growth, feed efficiency
Leucine, %
1.26
1–3
Growth, feed efficiency
Lysine, %
0.60
Fattening
Growth
0.90
0–8
Growth, Plasma lysine
0.64
3–6
Growth
0.55
6–10
Growth
1.06
1–3
Growth, feed efficiency
<0.70
1–7
Growth, feed efficiency
Methionine, %
0.45
0–1.5
Growth
0.30
3–6
Growth
Breed
References
White Pekin
White Pekin
White Pekin
White Pekin
White Pekin
White Pekin
Dean, 1972a
Dean, 1972a
Wilson, 1975
Wilson, 1975
Siregar et al., 1982
Siregar et al., 1982
Mule
Chen and Shen, 1979
Mule
Yu and Shen, 1984
Mule
Yu and Shen, 1984
Not specified
Pekin
Muscovy
Muscovy
Mule
Pekin
Jeroch and Hennig, 1965
Gazo et al., 1970
Leclerq and Carville,1977
Leclerq and Carville, 1977
Chen and Shen, 1979
Adams et al., 1983
Pekin
Muscovy
Dean, 1967
Leclerq and de Carville,
1977a
Leclerq and de Carville,
1977a
Elkin et al., 1986
0.25
6–10
Growth
Muscovy
0.40
Methionine + cystine, %
0.60
0.60
0–2
Growth
Pekin
0–1.5
3–6
Growth
Growth
Pekin
Muscovy
0.55
6–10
Growth
Muscovy
0.70
Tryptophan, %
0.23
Valine, %
0.78
Calcium, %
0.56
0–2
Growth
Pekin
Dean, 1967
Leclerq and de Carville,
1977a
Leclerq and de Carville,
1977a
Elkin et al., 1986
1–3
Growth, feed efficiency
Mule
Wu et al., 1984
1–3
Growth, feed efficiency
Mule
Yu and Shen, 1984
0–8
Pekin
Dean et al., 1967
Pekin
Taiwan
Taiwan
Dean, 1972b
Su, 1977
Su, 1977
0.58
1.00
3.75
Nonphytate phosphorus, %
0.60
1.05
0.40
Ducklings
Ducklings
Sexually mature
Growth, feed efficiency,
bone ash
Growth, bone ash
Growth
Egg production
0–4
Sexually mature
0–3
Growth, bone ash
Egg production
Growth
Pekin
Taiwan
Muscovy
0.22
3–6
Growth
Muscovy
0.18
6–10
Growth
Muscovy
0.34
Sodium chlorine, %
0.14
0.12
Magnesium, mg/kg
500
0–3
Growth, bone ash
Mule
Dean, 1972a
Su, 1977
Leclerq and de Carville,
1979
Leclerq and de Carville,
1979
Leclerq and de Carville,
1979
Lin and Shen, 1979
0–7
0–7
Growth, liveability
Growth, liveability
Pekin
Pekin
Dean, 1972a
Dean, 1972a
0–2
Growth, brain alkaline
phosphatase
Pekin
Van Reen and Pearson,
1953
0–3
Growth
Mule
Wu and Shen, 1978
Manganese, mg/kg
50
Zinc, mg/kg
68
Selenium, mg/kg
0.14
0–3
Growth
Mule
Wu and Shen, 1978
0–7
Pekin
Dean and Combs, 1981
0.20
0–7
Growth, liveability,
glutathione peroxidase
Growth liveability
glutathione peroxidase
Pekin
Dean and Combs, 1981
Vitamin D3, IU/kg
300
400
0–3
0–3
Bone ash
Bone ash
Pekin
Pekin and Indian
Runner
Fritz et al., 1941
Motzok and branion, 1946
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APPENDIXES
Nutrient and Estimated
Requirement
Vitamin E, IU/kg
9
Vitamin K, mg/kg
0.5
Riboflavin, mg/kg
3
4
Pantothenic acid, mg/kg
11
Niacin, mg/kg
52
45
Pyridoxine, mg/kg
2.5
108
Age Period (Days)
Response Criteria
Breed
References
0–4
Myopathy of heart muscle and
smooth muscle of intestines
Pekin
Jager, 1972
0–2
Prothrombin time
Pekin
Dean, 1972
0–7
0.5–2
Growth
Growth
Pekin
Pekin
Fritz et al., 1939
Hegsted and Perry, 1948
0.5–2
Growth
Pekin
Hegsted and Perry, 1948
0–2
0–3
Growth, leg development
Growth, feed efficiency
Pekin
Mule
Heuser and Scott, 1953
Wu et al., 1984
0.5–3 or longer
Growth, hemoglobin, hematocrit
Pekin
Hegsted and Rao, 1945
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use the print version of this publication as the authoritative version for attribution.
Nutrient Requirements of Poultry: Ninth Revised Edition, 1994
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APPENDIXES
109
TABLE A-10 Documentation of Nutrient Requirements of Pheasants
Nutrient and Estimated
Age Period (Days)
Response Criteria
Requirement
Metabolizable energy, kcal/kg
2,700
Sexually mature
Egg production, egg weight, feed
efficiency, mortality
Protein, %
26
0–3
Growth
24
3–5
Growth
26
0–4
Growth, feed efficiency
24
0–8
Growth, feathering, liveability
20
8 – 16
Growth, feathering, liveability
12
After 16
Growth, feathering, liveability
28
0–4
Growth
28
0–4
Growth, feed efficiency
19
8 – 17
Growth, feathering, feed
efficiency, liveability
15
Sexually mature
Egg production, fertility,
hatchability
Methionine, %
0.48
0–4
Growth
Methionine + cystine, %
0.94
0–4
Growth
Calcium, %
0.93
0–5
Growth, bone ash
0.53
5 – 14
Growth, bone ash
0.90
0–5
Growth, bone ash
1.2
0–8
Growth, bone ash
2.1
Sexually mature
Egg production, shell quality,
bone ash
2.0
Sexually mature
Egg production, fertility,
hatchability, body weight
Total phosphorus, %
0.98
0–4
Growth, bone ash
0.7
0–5
Growth, bone ash
0.48
5 – 14
Growth, bone ash
Nonphytate phosphorus, %
0.6
0–8
Growth, bone ash
0.6
Sexually mature
Egg production bone ash
Sodium, %
0.22
0–4
Growth, liveability
Manganese, mg/kg
70
0–5
Growth, bone development
Zinc, mg/kg
62
0–5
Growth, feather and bone
development
120
0–3
Growth, feather development
Vitamin D3, IU/kg
1,500
0–5
Growth, bone ash
Riboflavin, mg/kg
3.4
0–5
Growth, feather and bone
development
Pantothenic acid, mg/kg
10
0–4
Growth, feather and bone
development
Niacin, mg/kg
50
0–4
Growth, bone development
70
0–5
Growth, feathering and bone
development
Choline, mg/kg
0–5
Growth, feather and bone
1,430
development
Breed
References
Ring–neck
Monetti et al., 1982
Ring–neck
Ring–neck
Ring–neck
Chinese
Chinese
Chinese
Ring–neck
Ring–neck
Ring–neck
Scott et al., 1954
Scott et al., 1954
Scott et al., 1963
Woodard et al., 1977
Woodard et al., 1977
Woodard et al., 1977
Fuentes, 1981
Warner et al., 1982
Cain et al., 1984
Ring–neck
Monetti et al., 1985
Ring–neck
Fuentes, 1981
Ring–neck
Fuentes, 1981
Ring–neck
Ring–neck
Ring–neck
Ring–neck
Ring–neck
Scott et al., 1958a
Scott et al., 1958a
Hinkson et al., 1971
Reynnells, 1979
Reynnells, 1979
Ring–neck
Wise and Ewins, 1980
Ring–neck
Ring–neck
Ring–neck
Sunde and Bird, 1956
Scott et al., 1958a
Scott et al., 1958a
Ring–neck
Ring–neck
Reynnells, 1979
Reynnells, 1979
Ring–neck
Scott et al., 1960
Ring–neck
Scott et al., 1959
Ring–neck
Scott et al., 1959
Ring–neck
Cook et al., 1984
Ring–neck
Scott et al., 1958a
Ring–neck
Scott et al., 1959
Ring–neck
Scott et al., 1964
Ring–neck
Ring–neck
Sunde and Bird, 1957
Scott et al., 1959
Ring–neck
Scott et al., 1959
Copyright © National Academy of Sciences. All rights reserved.
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APPENDIXES
110
TABLE A-11 Documentation of Nutrient Requirements of Japanese Quail
Nutrient and Estimated
Age Period (Days)
Response Criteria
Requirement
Protein, %
24
0–35
Growth, protein retention
24
0–42
Growth
26
0–35
Growth, feed efficiency
25
0–28
Growth
20
Sexually mature
Egg production, egg weight,
feed efficiency
20
Sexually mature
Egg production
28.4
Sexually mature
Egg production
16
Sexually mature, peak egg
Egg production, egg yield, body
production
weight
24
Sexually mature
Not specified
20
Sexually mature
Egg production
24
0–28
Growth, carcass characteristics
Arginine, %
1.25
0–10
Growth
1.13
Sexually mature
Egg production, body weight,
egg weight
Glycine, %
1.74
0–21
Growth
1.17
21–35
Growth
Glycine + serine, %
1.14
0–10
Growth
Histidine, %
0.36
0–10
Growth
0.38
Sexually mature
Egg production, body weight,
egg weight
Isoleucine, %
0.98
0–10
Growth
0.81
Sexually mature
Egg production, body weight,
egg yield
Leucine, %
1.69
0–10
Growth
1.28
Sexually mature
Egg production, body weight,
egg weight
Lysine, %
1.37
0–21
Growth
1.2
21–35
Growth
1.15
0–10
Growth
0.86
Sexually mature
Egg production
0.97
Sexually mature
Egg production
Methionine, %
0.43
0–10
Growth
0.37
Sexually mature
Egg production, body weight,
egg yield
0.48
0–35
Growth, feed efficiency, feather
development, carcass yield
0.27
Sexually mature
Egg production
0.39
Sexually mature
Egg production, feather loss
Methionine + cystine, %
0.74
0–21
Growth
0.72
21–35
Growth
0.72
0–10
Growth
0.68
Sexually mature
Egg production, body weight,
egg yield
0.75
0–35
Growth, feed efficiency, feather
development, carcass yield
0.72
Sexually mature
Egg production
0.71
Sexually mature
Egg production, feather loss
Phenylalanine + tyrosine, %
1.79
0–10
Growth
1.25
Sexually mature
Egg production, body weight,
egg yield
Threonine, %
1.02
0–10
Growth
0.67
Sexually mature
Egg production, body weight,
egg yield
Tryptophan, %
0.22
0–10
Growth
0.17
Sexually mature
Egg production, body weight,
egg yield
Valine, %
0.95
0–10
Growth
Sexually mature
Egg production, body weight,
0.83
egg yield
Copyright © National Academy of Sciences. All rights reserved.
References
Weber and Reid, 1967
Lepore and Marks, 1971
Vogt, 1969
Vohra and Roudybush, 1971
Begin and Insko, 1972
Lee et al., 1977
Sakurai, 1979
Allen and Young, 1980
Sakurai, 1981
Shim and Lee, 1982
Steigner, 1990
Young et al., 1978
Allen and Young, 1980
Svacha et al., 1970
Svacha et al., 1970
Young et al., 1978
Young et al., 1978
Allen and Young, 1980
Young et al., 1978
Allen and Young, 1980
Young et al., 1978
Allen and Young, 1980
Svacha et al., 1970
Svacha et al., 1970
Young et al., 1978
Allen and Young, 1980
Shim and Lee, 1984
Young et al., 1978
Allen and Young, 1980
Shrivastav and Panda, 1987
Shim and Lee, 1988
Shim and Lee, 1989
Svacha et al., 1970
Svacha et al., 1970
Young et al., 1978
Allen and Young, 1980
Shrivastav and Panda, 1987
Shim and Lee, 1988
Shim and Chen, 1989
Young et al., 1978
Allen and Young, 1980
Young et al., 1978
Allen and Young, 1980
Young et al., 1978
Allen and Young, 1980
Young et al., 1978
Allen and Young, 1980
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APPENDIXES
111
Nutrient and Estimated
Requirement
Calcium, %
2.5
0.80
Age Period (Days)
Response Criteria
References
Sexually mature
0–14
Nelson et al., 1964
Consuegra and Anderson, 1967
0.48
14–28
0.44
0–35
0.70
Nonphytate phosphorus, %
0.6
0.30
0–21
Egg production, hatchability
Growth, bone ash, calcium and
phosphorus retention
Growth, bone ash, calcium and
phosphorus retention
Growth, feed efficiency, bone ash,
liveability
Growth, bone ash
Nelson et al., 1964
Consuegra and Anderson, 1967
0–21
Egg production, hatchability
Growth, bone ash, calcium and
phosphorus retention
Growth, bone ash
0–28
8–35
Growth
Growth, liveability, adrenal weight
Scott et al., 1960
Lumijarvi and Vohra, 1976
0–14
Growth, liveability, hemoglobin,
tibia ash
Growth, liveability
Harland et al., 1976
0.3
Sodium chlorine, %
0.15
0.10
Magnesium, mg/kg
300
150 mg
Iron, mg/kg
120
Sexually mature
0–28
0–14
Consuegra and Anderson, 1967
Miller, 1967
Bisoi et al., 1980
Bisoi et al., 1980
Vohra, 1972b
0–28
Growth, hemoglobin, feathering,
bone ash
Harland et al., 1973
Copper, mg/kg
<5
0–28
Growth, hemoglobin, feathering,
bone ash
Harland et al., 1973
Manganese, mg/kg
<12
0–28
Growth, hemoglobin, feathering,
bone ash
Harland et al., 1973
Zinc, mg/kg
25
0–28
Growth, feathering, tibia ash,
liveability
Spivey-Fox and Jacobs, 1967
0–42
Growth, liveability
Thompson and Scott, 1967
0–28
Growth, thyroid weight
Scott et al., 1960
7–56
Sexually mature
0–14
0–10
Sexually mature
Growth, liveability
Hatchability
Growth
Growth, liver vitamin A
Hatchability, liveability, vitamin A
in yolk
Shellenberger and Lee, 1966
Shellenberger and Lee, 1966
Ramachandran and Arscott, 1974
Parrish and Al-Hasani, 1983
Parrish and Al-Hasani, 1983
0–21
0–14
Bone ash, plasma calcium
Growth
Shue, 1967
Ramachandran and Arscott, 1974
0–14
0–35
Growth
Growth, liveability
Ramachandran and Arscott, 1974
Mak and Vohra, 1982
0–14
0–35
Growth
Growth, viability
Ramachandran and Arscott, 1974
Mak and Vohra, 1982
0–7
Spivey-Fox et al., 1966
Selenium, mg/kg
0.1
Iodine, mg/kg
0.3
Vitamin A, IU/kg
1,650
3,300
825
1,000
3,200
Vitamin D, IU/kg
480
750
Thiamine, mg/kg
6
1.2
Niacin, mg/kg
40
15
Pantothenic acid, mg/kg
40
10
7–35
10
15
23
Riboflavin, mg/kg
8
2
Choline, mg/kg
2,500
2,090
1,045–2,090
1,300
Folacin, mg/kg
0.36
Pyridoxine, mg/kg
6
1,25
0–35
Sexually mature
0–14
Growth, feather development,
dermatitis
Growth, feather development,
dermatitis
Growth, feather development
Fertility, hatchability
Growth
0–14
0–35
Growth
Growth, viability
Ramachandran and Arscott, 1974
Mak and Vohra, 1982
0–28
Sexually mature
Sexually mature
0–14
Growth, feed efficiency
Egg weight
Body weight, liver lipids
Growth
Vogt, 1970
Latshaw and Jensen, 1971
Latshaw and Jensen, 1972
Ramachandran and Arscott, 1974
Not specified
Growth, liveability
Wong et al., 1977
0–14
0–35
Growth
Growth, viability
Ramachandran and Arscott, 1974
Mak and Vohra, 1982
Spivey-Fox et al., 1966
Cutler and Vohra, 1967
Cutler and Vohra, 1967
Ramachandran and Arscott, 1974
Copyright © National Academy of Sciences. All rights reserved.
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Nutrient Requirements of Poultry: Ninth Revised Edition, 1994
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APPENDIXES
112
TABLE A-12 Documentation of Nutrient Requirements of Bobwhite Quail
Nutrient and Estimated Requirement
Age Period (Days)
Response Criteria
Metabolizable energy, kcal/kg
2,850–3,170
0–5
Growth, energy consumption, feed efficiency
Protein, %
28
0–8
Growth, liveability
20
0–6
Growth, liveability
26.5
0–4
Growth, feed efficiency, feathering
28
0–6
Growth
20
6–9
Growth
26
0–5
Growth, feed efficiency
24
0–5
Growth, feed efficiency
Methionine + cystine, %
1.0
0–5
Growth
Calcium, %
0.65
0–6
Growth, liveability, bone ash
2.3
Sexually mature
Egg production, eggshell thickness, fertility,
hatchability
2.4
Sexually mature
Egg production, eggshell thickness, fertility
Nonphytate phosphorus, %
0.8
Sexually mature
Egg production, fertility, hatchability,
liveability of offspring
0.40
0–6
Growth, liveability, tibia ash
0.28
6–12
Growth, liveability, bone ash
0.45
0–6
Growth, liveability, bone ash
0.35
0–6
Growth, bone ash
>0.70
Sexually mature
Egg production, egg shell thickness, fertility
Vitamin A, IU/kg
8,800
0–10
Growth, liveability
13,200
Sexually mature
Reproduction, survival of offspring
Riboflavin, mg/kg
3.8
0–5
Growth, feed efficiency, liveability
Pantothenic acid, mg/kg
10
0–4
Growth, liveability, feathering, leg development
12.6
0–5
Growth, feed efficiency, liveability
Niacin, mg/kg
31
0–5
Growth, feed efficiency, liveability
Choline, mg/kg
0–5
Growth, feed efficiency, liveability
1,500
Copyright © National Academy of Sciences. All rights reserved.
References
Wilson et al., 1977
Baldini et al., 1950
Baldini et al., 1953
Scott et al., 1963
Andrews et al., 1973
Andrews et al., 1973
Serafin, 1977
Serafin, 1982
Serafin, 1982
Wilson et al., 1972
Dewitt et al., 1949
Cain et al., 1982
Dewitt et al., 1949
Scott et al., 1958b
Scott et al., 1958b
Wilson et al., 1972
Powell et al., 1974
Cain et al., 1982
Nestler, 1946
Nestler, 1946
Serafin, 1974
Scott et al., 1964
Serafin, 1974
Serafin, 1974
Serafin, 1974
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Nutrient Requirements of Poultry: Ninth Revised Edition, 1994
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APPENDIXES
113
TABLE B–1 Estimating the Energy Value (kcal/kg dry matter) of Feed Ingredients from Proximate Composition (components as
percentage of ingredient unless otherwise noted)
Ingredient
Prediction Equation
Reference
Cereal grains and milling by-products
Corn grain
MEn = 36.21 × CP + 85.44 × EE + 37.26 × NFE
Janssen, 1989
Sorghum (tannin <0.4%)
MEn = 31.02 × CP + 77.03 × EE + 37.67 × NFE
Janssen, 1989
Janssen, 1989
Sorghum (tannin >1.0%)
MEn = 21.98 × CP + 54.75 × EE + 35.18 × NFE
Sorghum
ME = 3,152 - 357.79 × tannic acid
Gous et al., 1982
Janssen, 1989
Sorghum
MEn = 38.55 × DM - 394.59 × tannic acid
Sorghum
ME = 3,062 + 887 × CF - 202.5 × (CF)2
Moir and Connor, 1977
Sorghum
ME = 4,412 - 90.34 × ADF
Moir and Connor, 1977
2
Moir and Connor, 1977
Sorghum
ME = 3,773 + 65.73 × APF - 3.272 × (APF)
Janssen, 1989
Triticale
MEn = 34.49 × CP + 62.16 × EE + 35.61 × NFE
Janssen, 1989
Wheat
MEn = 34.92 × CP + 63.1 × EE + 36.42 × NFE
Janssen, 1989
Polished rice, rice polishings
MEn = 46.7 × DM - 46.7 × ash - 69.55 × CP +
42.95 × EE - 81.95 × CF
Janssen, 1989
Rice bran, solvent extracted
MEn = 46.7 × DM - 46.7 × ash - 69.54 × CP +
42.94 × EE - 81.95 × CF
Rice products
MEn = 4,759 - 88.6 × CP - 127.7 × CF + 52.1 × EE
Janssen et al., 1979
Janssen, 1989
Bakery by-product
MEn = 34.49 × CP + 76.1 × EE + 37.67 × NFE
Dale et al., 1990
Dried bakery products
TMEn = 4,340 - 100 × CF - 40 × ash - 30 × CP +
10 × EE
Janssen, 1989
Wheat middlings, wheat bran
MEn = 40.1 × DM - 40.1 × ash - 165.39 × CF
Janssen et al., 1979
Wheat and wheat products (feeds in meal form)
MEn = 3,985 - 205 × CF
Janssen et al., 1979
Wheat and wheat products (feeds in pellet form)
MEn = 3,926 - 181 × CF
Janssen et al., 1979
Barley and barley products
MEn = 3,078 - 90.4 × CF + 9.2 × STA
Janssen et al., 1979
Oats and oat products
MEn = 2,970 - 59.7 × CF + 116.9 × EE
Starch industry by-products
Corn wet-milling by-products
MEn = 4,240 - 34.4 × CP - 159.6 × CF + 13.5 × EE
Janssen et al., 1979
Janssen, 1989
Corn gluten meal (65% crude protein)
MEn = 40.94 × CP + 88.17 × EE + 33.13 × NFE
Janssen, 1989
Corn gluten meal (40% crude protein)
MEn = 36.64 × CP + 73.3 × EE + 25.67 × NFE
Janssen, 1989
Corn gluten feed (20% crude protein)
MEn = 42.35 × DM - 42.35 × ash - 23.74 × CP +
28.03 × EE - 165.72 × CF
Sugar industry products
Beet or cane molasses
MEn = 40.01 × SUG
Janssen, 1989
Janssen, 1989
Sugar
MEn = 38.96 × SUG
Distillers by-products
Janssen, 1989
Brewer's dried grains, corn distillers' dried solubles,
MEn = 39.15 × DM - 39.15 × ash - 9.72 × CP 63.81 × CF
corn distillers' dried grains, corn distillers' dried
grains plus solubles
Janssen, 1989
Yeast, torula
MEn = 34.06 × CP + 40.82 × EE + 26.91 × NFE
Dried roots
Janssen, 1989
Sweet potatoes (dried)
MEn = 8.62 × CP + 50.12 × EE + 37.67 × NFE
Janssen, 1989
Tapioca meal (e.g., cassava)
MEn = 39.14 × DM - 39.14 × ash - 82.78 × CF
Janssen et al., 1979
Tapioca meal (e.g., cassava)
MEn = 4,054 - 43.4 × ash - 103 × CF
Oilseeds, oilseed meals, and by-products
Cottonseed meal, expeller or solvent
MEn = 21.26 × DM + 47.13 × EE - 30.85 × CF
Janssen, 1989
Janssen et al., 1979
Cottonseed products
MEn = 2,153 - 31.8 × CF + 43.5 × EE
Janssen, 1989
Peanut meal, expeller or solvent
MEn = 29.68 × DM + 60.95 × EE - 60.87 × CF
Janssen et al., 1979
Peanut products
MEn = 3,072 - 39.1 × ash - 47.6 × CF + 63.7 × EE
Janssen, 1989
Rapeseed meal, solvent, high glucose
MEn = 29.73 × CP + 46.39 × EE + 7.87 × NFE
Rapeseed meal, solvent, double zero
MEn = 32.76 × CP + 64.96 × EE + 13.24 × NFE
Janssen, 1989
Janssen, 1989
Soybean meal, expeller
MEn = 37.5 × CP + 70.52 × EE + 14.9 × NFE
Janssen, 1989
Soybean meal, solvent
MEn = 37.5 × CP + 46.39 × EE + 14.9 × NFE
Janssen et al., 1979
Soybean meal (solvent or expeller process)
MEn = 2,702 - 57.4 × CF + 72.0 × EE
Janssen, 1989
Soybeans, heat treated, meal
MEn = 36.63 × CP + 77.96 × EE + 19.87 × NFE
Janssen, 1989
Soybeans, heat treated, pellet
MEn = 38.79 × CP + 87.24 × EE + 18.22 × NFE
Full-fat soybeans (feeds in meal form)
MEn = 2,769 - 59.1 × CF + 62.1 × EE
Janssen et al., 1979
Janssen et al., 1979
Full-fat soybeans (feeds in pellet form)
MEn = 2,636 - 55.7 × CF + 82.5 × EE
Sunflower seeds, unextracted
MEn = 36.64 × CP + 89.07 × EE + 4.97 × NFE
Janssen, 1989
Janssen et al., 1979
Sunflower products
MEn = 3,999 - 189 × ash - 58.5 × CF + 59.5 × EE
Janssen, 1989
Sunflower, expeller, with hulls
MEn = 26.7 × DM + 77.2 × EE - 51.22 × CF
Sunflower, expeller or solvent, decorticated
MEn = 6.28 × DM - 6.28 × ash + 25.38 × CP 62.62
Janssen, 1989
× EE
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Nutrient Requirements of Poultry: Ninth Revised Edition, 1994
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APPENDIXES
114
Ingredient
Products of animal origin
Skim milk powder
Whey, dried, low lactose
Meat and bone meal
Fish meal (60%, 65%, 67% crude protein)
Herring meal, Norwegian
Blood meal, spray dried
Blood meal, drum dried
Feather meal (pepsin dig 80%)
Poultry offal meal
Poultry offal meal, high-fat
Poultry by-product meal
Poultry by-product meal
Poultry by-product meal
Poultry by-product meal
Poultry by-product meal
Poultry by-product meal
Poultry by-product meal
Poultry by-product meal
Poultry by-product meal
Fat products from Dutch renderers
Fats and oils
All fats and oils
Vegetable oils (free fatty acid <50%)
Vegetable oils (free fatty acid >50%)
Animal fats (free fatty acid <40%)
Animal fats (free fatty acid >40%)
Prediction Equation
Reference
MEn = 40.94 × CP + 77.96 × EE + 19.04 × NFE
MEn = 38.79 × CP + 77.96 × EE + 19.04 × NFE
MEn = 33.94 × DM = 45.77 × ash + 59.99 × EE
MEn = 35.87 × DM - 34.08 × ash + 42.09 × EE
MEn = 35.87 × DM - 34.08 × ash + 42.09 × EE
MEn = 34.49 × CP + 64.96 × EE
MEn = 31.88 × CP + 60.32 × EE
MEn = 33.2 × CP + 57.53 × EE
MEn = 31.02 × CP + 74.23 × EE
MEn = 31.02 × CP + 78.87 × EE
TMEn = -725 + 0.841 × GE (kcal/kg dry matter)
TMEn = 4,070 - 142 × calcium
TMEn = 4,330 - 61 × ash
TMEn = 5,060 - 263 × ash + 491 × calcium
TMEn = 479 + 89 × CP - 1,094 × phosphorus
TMEn = 11,340 - 103 × CP - 327 × calcium
TMEn = 934 - 69 × CP - 110 × ash
TMEn = 561 - 154 × calcium - 622 × phosphorus
TMEn = 556 - 63 × ash - 506 × phosphorus
MEn = 20,041 - 23.0 × IV - 319.1 × C16 : 0 - 153.4 × C18 : 0
MEn = 8,227 - 10,318(-1,1685[Unsaturated:Saturated ratio])
MEn = 28,119 - 235.8 (C18 : 1 + C18 : 2) - 6.4 (C16:0) 310.9 (C18 : 0) + 0.726 (IV × FR1) - 0.0000379 (IV[FR1 +
FFA])2
MEn = -10,147.94 + 188.28 IV + 155.09 FR1 - 1.6709 (IV
× FR1)
MEn = 1,804 + 29.7084 IV + 29.302 FR1
MEn = 126,694 + 1645 IV + 838.4 C16 : 0 - 215.3 C18 : 0
+ 746.61 FR1 + 356.12 (FR1 + FFA) - 14.83 (IV × FR1)
MEn = -9,865 + 194.1 IV + 300.1 C18:0
Janssen, 1989
Janssen, 1989
Janssen, 1989
Janssen, 1989
Janssen, 1989
Janssen, 1989
Janssen, 1989
Janssen, 1989
Janssen, 1989
Janssen, 1989
Pesti et al., 1986
Pesti et al., 1986
Pesti et al., 1986
Pesti et al., 1986
Pesti et al., 1986
Pesti et al., 1986
Pesti et al., 1986
Pesti et al., 1986
Pesti et al., 1986
Janssen et al., 1979
Ketels and DeGroote, 1989
Huyghebaert et al., 1988
Huyghebaert et al., 1988
Huyghebaert et al., 1988
Huyghebaert et al., 1988
Huyghebaert et al., 1988
NOTE: Abbreviations used above are as follows: GE = gross energy; ME = metabolizable energy; MEn = nitrogen-corrected
metabolizable energy; TMEn = nitrogen-corrected true metabolizable energy; CP = % crude protein; EE = % ether extract; CF = % crude
fiber; NFE = % nitrogen-free extract; ADF = % acid detergent fiber; APF = % Acid-pepsin fiber; STA = % starch; SUG = % sugar; IV =
iodine value; C16 : 0 = % palmitic acid; C18 : 0 = % stearic acid; C18:1 = % oleic acid; C18 : 2 = % linoleic acid; FFA = % free fatty
acid, calculated as oleic acid equivalents; FR1 = first fraction from a column chromatography separation that contains the practically
unaltered triglycerides plus other apolar components; and DM = dry matter.
TABLE C–1 Conversion reactors—Weights and Measures
Units
Multiplied by the Factor Below
Equals
lb
453.6
lb
0.4536
oz
28.35
kg
1,000
kg
1,000,000
g
1,000
g
1,000,000
g
109
g
1012
mg
1,000
mg/kga
0.0001
ppm
0.0001
gal (U.S.)
3.785
4.546
liters
(hectoliter)
2.837
cal (calorie)
4.184
kcal (kilocalorie)
1,000
Mcal (megacalorie)
1,000,000
1,000
Mcal
a
Units
g
kg
g
g
mg
mg
mcg (or µg)
ng (nanogram)
pg (picogram)
mcg (or µg)
%
%
liters
0.220
bu
j (joule)
cal
cal
kcal
Multiplied by the Factor Below
Equals
0.002205
2.205
0.035273
0.001
0.000001
0.001
0.001
10-9
10-12
0.001
10,000
10,000
0.2642
gal (Brit.bu (bushel)
lb
lb
oz
kg
kg
g
mg
g
g
mg
mg/kg
ppm
gal (U.S.gal (Brit.)
0.3525 hl
0.239
0.001
0.000001
0.001
cal
kcal
Mcal
Mcal
100 ppm = 100 mg/kg = 0.010 percent; thus converting 0.0002 percent = 2 ppm = 2 mg/kg.
Copyright © National Academy of Sciences. All rights reserved.
Units
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AUTHORS
143
Authors
F. Howard Kratzer is professor emeritus of avian science at the University of California at Davis, from where he
received his Ph.D. in animal nutrition. His many concurrent positions include a visiting professorship at the University of
Sydney (Australia), and the Federal University of Rio Grande do Sol (Brazil). His research interests are poultry nutrition,
amino acid requirements of chickens and turkeys, vitamin needs and functions, minerals and mineral availability, and growth
inhibitors.
J. David Latshaw is professor of poultry science at Ohio State University, where he has taught since 1970. He received
his Ph.D. in nutrition from Washington State University. Research areas of major interest to him are factors influencing feed
intake in poultry, and interaction of diet and growth efficiency.
Steven L. Leeson currently is a professor of poultry science at the University of Guelph (Canada). He received his
Ph.D. in poultry nutrition from the University of Nottingham (England). His research areas are feeding programs for leghorn
birds, interaction of nutrient supply from feed and body reserves, and energy evaluation of ingredients.
Edwin T. Moran, Jr., previously a professor at the University of Guelph (Canada), Moran has been professor of animal
nutrition at Auburn University since 1986. He received his Ph.D. in animal nutrition from Washington State University. His
research experience includes influence of nutrition and management on broiler yields, amino acid availability and
performance, and feedstuff evaluation in broiler production.
Carl M. Parsons currently is assistant professor of animal science at the University of Illinois at Urbana-Champaign.
He received his Ph.D. in animal science from Virginia Polytech Institute and State University. Research interests include
poultry production and management with emphasis in the field of nutrition, and improved nutritional efficiency for
production of poultry meat and eggs, particularly with respect to protein utilization.
Jerry L. Sell (Chair) is professor of animal nutrition at Iowa State University, where he has taught since 1976, and from
where he received his Ph.D. in poultry nutrition. His major areas of research are energy efficiency of chickens and turkeys,
and metabolism of minerals.
Park W. Waldroup is professor of poultry nutrition at the University of Arkansas at Fayetteville. He received his Ph.D.
in nutritional biochemistry from the University of Florida. Among his research interests are studies concerned with nutrient
requirements of poultry in terms of nutrient balance and interrelationships of nutrients, and effects of processing on nutritive
value of feed.
Copyright © National Academy of Sciences. All rights reserved.
AUTHORS
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144
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INDEX
145
Index
A
Absorption
in calcium metabolism, 50
high level dietary fat and, 12
of vitamin K, 52
of xanthophylls, 17
Acid-base balance, 14
Aflatoxins, 78-79
Age
broiler nutrient requirements and, 27
fat metabolism and, 12
metabolizable energy assessment and, 6
Alfalfa
amino acid composition of, 66, 71
amino acid digestibility coefficients, 74
fatty acid composition of, 75
nutrient composition of, 72-63
xanthophylls in, 17
Aluminum, toxicity of, 59
Amaranth, 53
Amino acids
biochemical role of, 9
in broiler breeder diet, 32-33, 34
in broiler chicken diet, 27-29
conversion to vitamins, 11
crystalline form, 28, 80
deficiencies in chickens and turkeys, 46-50
digestibility, 71, 73-74
documentation of nutrient requirements, 85
duck requirements, 43
egg weight and, 25
in feeds, 61, 66-68, 69, 71-74
geese requirements, 40-41
interactions, 10-11
metabolizable energy of, 75
in nutritional requirement models, 1
in phase-feeding program, 24
requirements, 9-10
supplements, 74
total sulfur amino acids, 28, 38, 41, 43, 45
toxicity, 11
turkey requirements, 36, 37-38
Animal products
as dietary fat source, 11, 12
nutrient composition of meat, 72-63, 67
as protein source, 70
Antibiotic additives, 18
Antimicrobial additives, 3, 18, 52
Antioxidants, 50, 51
Arginine, 11
broiler chicken requirements, 29
deficiency, 46
documentation of nutrient requirements, 90, 99, 107, 110
duck requirements, 43
in feeds, 66-68, 71-73
turkey requirements, 37-38
Arsenic, toxicity of, 59
B
Bakery waste products
amino acid composition, 66, 72
amino acid digestibility coefficients, 74
metabolizable energy in, 6, 113
nutrient composition of, 72-63
Barium, toxicity of, 59
Barley, 8, 9
amino acid composition of, 71, 73
amino acid digestibility coefficients, 74
fatty acid composition of, 75
metabolizable energy in, 113
nutrient composition of, 72-63
Bicarbonate, 14
in egg production, 25
Bioassay techniques for measurement of energy values, 4-6
Biotin, 15
biochemical role of, 53
broiler breeder chicken requirements, 33
broiler chicken requirements, 31
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146
deficiency, 47, 48, 53-54
documentation of nutrient requirements, 87, 89, 95, 98, 105
in feeds, 63, 65
in wheat diet, 69
Blood meal
amino acid digestibility coefficients, 74
nutrient composition of, 72-63, 66
Bobwhite quail, 45, 112
Bone development
in Leghorn-type pullet, 21
mineral requirements, 14
nutrient deficiencies and, 49, 51, 53, 54, 55, 56, 57
vitamin D in, 50-51
Bone meal
amino acid composition of, 72
element concentrations in, 78
Boron, 15
Breeding diets
biotin in, 54
broiler chickens, 32-34
iodine in, 57
Japanese quail, 45
Leghorn-type laying chickens, 26
turkeys, 38, 39
zinc in, 57
Brewer's grains, nutrient composition of, 72-63, 66, 113
Broadbeans, nutrient composition of, 72-63, 66
Broiler breeder chickens
amino acid/protein requirements, 32-33, 34
documentation of nutrient requirements, 97-98
energy consumption, 33, 34
feed intake, 32
hens, 32-33
males, 33-34
mineral requirements, 33, 34
nutrient requirements, 32
vitamin requirements, 33
Broiler chickens
age of, and nutrient requirements, 27
amino acid requirements, 27-29
documentation of nutrient requirements, 90-96
energy intake, 8
fat utilization, 7
fatty acid requirements, 32
feed intake, 26
feed utilization vs. weight gain, 26-27
gender differences, 27
market weights, 26
mineral requirements, 27, 29-31
nutrient interactions in, 27
nutritional modeling for, 1
protein requirements, 27
vitamin requirements, 27, 31-32
water intake, 16
Buckwheat, nutrient composition of, 72-63, 66
Bushel weights of grains, 61, 68
C
Cadmium, toxicity of, 59
Calcium
biochemical role of, 14
Bobwhite quail requirements, 45
broiler breeder chicken requirements, 33, 34
broiler chicken requirements, 29, 30
deficiency, 47, 56
documentation of nutrient requirements, 85, 88, 93, 98,
102-103, 105, 106, 107, 109, 111, 112
duck requirements, 43
excesses of, 14
in feeds, 72, 64, 78
geese requirements, 41
in laying hen diet, 25
in Leghorn-type pullet growth, 22
in phase-feeding program, 24
toxicity to pheasant chicks, 44
turkey requirements, 38
vitamin D and, 50, 51
Calcium carbonate, 78
Calcium phosphate, 78
Calcium sulfate, 78
Calorie, definition of, 3
Canola
amino acid composition of, 71
amino acid digestibility coefficients, 74
nutrient composition of, 72-63, 66, 70
Carbohydrates
metabolizable energy values of, 75, 76-77
sources of, 8-9
β-Carotene
as pigmenting agent, 17
as vitamin A, 15, 50
Carotenoids
for pigmentation, 17, 50
Casein
amino acid digestibility coefficients, 74
nutrient composition of, 72-63, 66
Cereal grains
bushel weight ranges, 61, 68
as carbohydrate source, 8-9
metabolizable energy of, 61, 72, 64, 113
nutrient composition of, 61-69
Chickens
standard reference diets for, 80-81
See also Broiler chickens;
Broiler breeder chickens;
Leghorn-type
chickens;
Symptoms of nutrient deficiency
Chloride
biochemical role of, 14
in egg production, 25
toxicity, 59
Chlorine
broiler breeder chicken requirements, 33
deficiency, 47, 57
documentation of nutrient requirements, 86, 88, 94, 98, 103
in feeds, 72, 64, 78
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Choline, 11
broiler chicken requirements, 30, 31
chloride, 15
deficiency, 10, 47, 56
documentation of nutrient requirements, 87, 89, 95, 104,
105, 106, 109, 111, 112
in egg production, 25
in feeds, 63, 65
geese requirements, 41
in methionine-cystine requirements, 28
role of, 55
turkey requirements, 39
Chromium, toxicity of, 59
Cobalt, 14
toxicity, 59
Coconut
amino acid digestibility coefficients, 74
nutrient composition of, 72-63, 66
Copper, 14
biochemical role of, 57
broiler chicken requirements, 30
deficiency, 47, 48, 57
documentation of nutrient requirements, 86, 89, 94, 111
in feeds, 63, 65, 78
toxicity, 58, 59
Coprophagy, 52, 54
Corn
amino acid composition of, 71, 73
amino acid digestibility coefficients, 74
fatty acid composition of, 75
metabolizable energy in, 113
nutrient composition of, 72-63, 66
xanthophylls in, 17
Cost factors
in egg weight gain, 25
protein sources, 10
in setting dietary energy levels, 7
Cotton/cottonseed meal
amino acid composition of, 71
amino acid digestibility coefficients, 74
fatty acid composition of, 75
gossypol in, 69
metabolizable energy in, 113
nutrient composition of, 72-63, 66
as protein supplement, 69
Cystine
in broiler chicken diet, 27-28
documentation of nutrient requirements, 85, 88, 91-92,
101, 106, 107, 109, 110, 112
in feeds, 66-68, 71-73
methionine interaction, 10, 27-28
D
Deficiency, nutrient, 2
amino acid/protein, 10-11, 46-50
assessment of, 46, 47
embryo development and, 46
fatty acid, 13
feed consumption and, 7-8
to induce molt, 26
minerals, 47, 48, 56-57
trace element, 14
vitamins, 47, 48, 50-56
water deprivation, 16
See also Symptoms of nutrient deficiency
Dextrose, 80
Diagnosis of nutrient deficiency, 46
biochemical and physiological measurement in, 47
in embryo development, 46
Digestibility
of amino acids, 71, 73-74
measurement techniques, 73-74
of phosphorus, 14
of sorghums, 6
Digestible energy, 4
Drunken syndrome, 16
Ducks
documentation of nutrient requirements, 107-108
growing systems for, 42
nutrient requirements, 42-43
E
Egg production
in broiler breeder chickens, 32-33
calcium metabolism in, 56
carotenoid pigments in, 17
chloride/chlorine in, 25, 33
dietary fat in, 12-13
dietary minerals in, 14, 25
documentation of Leghorn-type chicken requirements,
88-89
linoleic acid in, 13
manganese in, 56
in pheasants, 44
phosphorus in, 33
potassium in, 56-57
pyridoxine in, 55
rapeseed meal diets and, 70
seasonal variation, 24
shell strength, 25, 33, 56, 57
sodium in, 33, 57
specific gravity of egg in, 33
in turkeys, 37, 38
vitamin D deficiency in, 51
yolk discoloration, gossypol in, 69
See also Embryo development;
Laying hens;
Leghorn-type chickens
Egg white, as protein source, 30
Eicosanoids, 13
Electrolytes, 14
Embryo development
biotin in, 54
choline in, 56
chorioallantois development in, 46
folic acid in, 55
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INDEX
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iodine in, 57
manganese in, 56
pantothenic acid in, 54
riboflavin in, 53
symptoms of nutrient deficiency in, 46
thiamin deficiency in, 53
vitamin B12 in, 55
vitamin D in, 51
vitamin K in, 52
Encephalomalacia, 31
Energy.
See also Metabolizable energy
carbohydrate sources, 8-9
digestible, 4
disposition of, 4
fat as source of, 11
gross, 4
net, 4
in setting dietary levels, 7-8
terminology, 3-4
Energy requirements
broiler breeder chickens, 33, 34
broiler chickens, 8
geese, 40
Leghorn-type laying hens, 22, 24
Leghorn-type pullets, 21
modeling of, 1
turkeys, 37
Established requirements, 1
Estimated requirements, 1
Ether extract, in feeds, 72, 64, 75
F
Fats
blending of, 12
depot fat, 13
dietary role of, 11
in egg weight, 25
energy values of, 6-7, 12-13, 114
in net energy of production, 12-13
saturated:unsaturated ratio, 7
sources of, 11-12
Fatty acids
broiler chicken requirements, 27, 32
composition in fats, 13
deficiencies, 13
dietary irregularities, symptoms of, 54
dietary synergism, 5
essential, 13
in feeds, 75-77
polyunsaturated, 13, 51-52
Feathers
in feeds, 72-63, 70, 74
growth in pheasants, 44
nutrient deficiencies in abnormalities of, 44, 46, 49, 50,
51, 52, 53, 54, 55, 57
Feces
in measuring energy, 4
phosphorus levels, 25
Feed manufacture/processing
amino acid availability and, 73
animal products in, 70
heat treatment of soybean meal, 70
mycotoxin formation, 78
pelleted feeds, 15-16, 35, 42, 53
Feedstuffs
amino acid composition of, 66-68, 71-74
analysis of, 2
antimicrobials in, 18
assessing energy in, 3-8
energy values of, prediction equations for, 113-114
fatty acid composition of, 75, 76-77
intake as factor in nutrient concentration in, 22-24
mineral concentrations in, 75-78
mycotoxins in, 78-79
nutrient composition of, 61-65, 68-70
pigmenting agents in, 17, 50
pyridoxine in, 54
thiamin in, 52-53
vitamin D in, 51
water:feed ratio, 15-16
Fermentation by-products, 18
Field beans
amino acid composition of, 71
amino acid digestibility coefficients, 74
Fish products, 53
amino acid composition of, 71, 73
amino acid digestibility coefficients, 74
metabolizable energy in, 114
nutrient composition of, 72-63, 66-67
as protein source, 70
Fluorine
in feeds, 78
toxicity, 59
Folacin/folic acid
antagonists, 55
broiler chicken requirements, 31
deficiency, 47, 48, 55
documentation of nutrient requirements, 87, 89, 95-96,
104, 105, 106, 111
in feeds, 63, 65
maximum tolerance, 15
metabolism, 55
Food and Drug Administration, 18
Formula diets, standard reference, 80-81
Fungi in feeds, 78-79
G
Geese
documentation of nutrient requirements, 106
force-feeding, 40
nutrient requirements, 40-41
protein requirements, 40
Gelatin
amino acid digestibility coefficients, 74
nutrient composition of, 72-63, 67
Gizzerosine, 70
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INDEX
149
Glucose
deficiency, 54
energy value, 4-5
Glycine
broiler chicken requirements, 29
documentation of nutrient requirements, 90, 99, 110
in feeds, 66-68
serine interaction, 10
Gossypol, 69
Gross energy, 4
Growth factors, unidentified, 3, 18
Growth/weight gain
amino acids and, 11
antimicrobials and, 18
in broiler breeder chickens, 32, 34
egg size and, 32-33
egg weight and, 24
in Leghorn-type chickens, 24
in Leghorn-type pullets, 19-21, 22
in market broilers, 26-27
in measuring metabolizable energy, 6
mineral deficiency and, 56-57
mineral toxicity and, 58
in turkey breeders, 39
in turkeys, 35
H
High-energy diets, 8
Histidine, 11
broiler chicken requirements, 29
deficiency, 47
documentation of nutrient requirements, 90, 97, 99, 110
in feeds, 66-68
Hominy, nutrient composition of, 72-63, 67
I
Iodine, 14
biochemical role of, 57
broiler chicken requirements, 31
deficiency, 47, 48, 57
documentation of nutrient requirements, 86, 89, 94, 111
to induce molt, 26
toxicity, 59
Iron, 14
broiler chicken requirements, 29, 30
deficiency, 47, 48, 57
documentation of nutrient requirements, 86, 88, 94, 111
in feeds, 63, 65, 78
pigmentation and, 57
toxicity, 59
Isoleucine, 11
broiler chicken requirements, 29
deficiency, 46, 50
documentation of nutrient requirements, 85, 88, 90, 100, 110
duck requirements, 43
in feeds, 66-68
leucine antagonism, 11
turkey requirements, 38
J
Japanese quail, 45, 110-111
Joule, definition of, 3-4
K
Kilocalorie, definition of, 3
L
Laying hens
added dietary fat for, 12-13
disposition of energy in, 4
energy intake, 8
nutritional modeling for, 1
vitamin D deficiency in, 51
water intake, 16
See also Egg production;
Leghorn-type chickens
Lead, toxicity of, 59
Leghorn-type chickens
ambient temperature in feed intake of, 8, 22
brown-egg-laying, 21, 25-26, 70
documentation of nutrient requirements of, 85-90
egg-type breeders, 26
egg weight, factors in, 24-25
energy requirements, 19-21, 22, 24
feed intake levels, 22-24
maintenance needs, 22
mineral requirements, 20, 21, 22, 25
molting hens, 26
phase feeding, 24
prelay period, 21-22
production diet, 22-26
protein requirements, 19-21, 22-24
pullet nutrient requirements, 19-22
vitamin requirements, 20, 21, 22, 25
Leucine, 11
broiler chicken requirements, 29
deficiency, 46, 50
documentation of nutrient requirements, 85, 90, 97, 100,
107, 110
duck requirements, 43
in feeds, 66-68
isoleucine antagonism, 11
Lighting, to induce molt, 26
Limestone, element concentrations in, 78
Linoleic acid, 13
broiler chicken requirements, 32
deficiency, 47
documentation of nutrient requirements, 93, 102, 105
egg weight and, 24
in feeds, 72, 64
Low-protein diets, 19-21
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150
Lupine seeds
amino acid composition of, 71, 72
amino acid digestibility coefficients, 74
Lutein, 17
Lysine, 11
in broiler chicken diets, 26-27, 29
deficiency, 46, 47
documentation of nutrient requirements, 85, 88, 90-91,
100, 106, 107, 110
duck requirements, 43
in feeds, 71-73
geese requirements, 40-41
Leghorn-type chicken requirements, 22-24
quail requirements, 45
turkey requirements, 37, 38
M
Magnesium
biochemical role of, 14
broiler chicken requirements, 29, 30
in calcium source, 14
deficiency, 47, 56
documentation of nutrient requirements, 86, 88, 94, 103,
107, 111
in feeds, 63, 65, 78
toxicity, 59
turkey requirements, 38
Magnesium oxide, element concentrations in, 78
Manganese, 14
broiler chicken requirements, 29, 30
deficiency, 47, 48, 56
documentation of nutrient requirements, 86, 88, 103, 105,
109, 111
in feeds, 63, 65, 78
toxicity, 59
turkey requirements, 38
Meat production
fish meal in feeds and, 70
See also Broiler chickens;
Broiler breeder chickens;
Ducks;
Geese;
Pheasants;
Quail;
Turkeys
Meat products as feed
amino acid composition of, 71, 72
amino acid digestibility coefficients, 74
element concentrations in, 78
fatty acid composition of, 75
fish products, 37, 53, 72-63, 66-67, 70, 71, 74
metabolizable energy in, 114
nutrient composition of, 72-63, 67
Megacalorie, definition of, 3
Menadione, 15
source of, 52
Menaquinone, 15
source of, 52
Mercury, toxicity of, 58, 59
Metabolizable energy
apparent, 4
carbohydrate values, 75, 76-77
of cereal grains, 68-69
definition of, 4
dietary requirements, 8
documentation of nutrient requirements, 98, 109, 112
in duck growth, 42
of fats, 12
of feed ingredients, 61, 72, 64
in geese, 40
measurement of, 4-6
prediction equations for estimating feed ingredient values,
113-114
proximate components in estimation of, 6-7
true, 4, 5, 6, 8
turkey feed intake and, 37
Methionine, 11
in broiler chicken diet, 27-28
cystine interaction, 10
deficiency, 50
documentation of nutrient requirements, 85, 88, 91, 97,
100-101, 106, 107, 109, 110, 112
duck requirements, 43
egg weight and, 24, 25
in feeds, 66-68, 71-73
geese requirements, 41
toxicity, 11
turkey requirements, 38
Milk, nutrient composition of, 72-63, 66
Millet, nutrient composition of, 72-63, 67
Milo, amino acid composition of, 71, 72
Minerals, 13-14
biochemical role of, 14
broiler breeder chicken requirements, 33, 34
broiler chicken requirements, 27, 29-31
deficiencies, 47, 48, 56-57
documentation of nutrient requirements, 85-86, 88-89,
93-94, 97, 98, 102-103, 105, 107, 109, 111, 112
duck requirements, 43
electrolyte balance, 14
in experimental diets, 15
in feeds, 78
interactions, 14-15, 56, 58
Leghorn-type chickens, requirements for, 20, 21, 22, 25, 26
macromineral supplements, 75-78
pheasant requirements, 44
supplements, 14-15
toxicity, 58-60
trace elements, 14
turkey requirements, 38
in water, 16-17
Modeling techniques
for amino acid requirements, 1
for energy requirements, 1
Molasses, nutrient composition of, 72-63
Molting hens, 26
Molybdenum, toxicity of, 59
Mycotoxins, 17
in feeds, 78-79
in vitamin D metabolism, 51
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INDEX
151
N
Net energy, definition of, 4
Niacin
amino acids in synthesis of, 11
biochemical role of, 53
broiler chicken requirements, 31
deficiency, 47, 48, 53
documentation of nutrient requirements, 87, 89, 95, 104,
105, 106, 108, 109, 111, 112
in feeds, 63, 65
geese requirements, 41
maximum tolerance, 15
Nickel, toxicity of, 59-60
Nitrate, toxicity of, 60
Nitrogen
in amino acids, 75
in measurement of metabolizable energy, 4, 5-6
Nutrient requirements
amino acid, 9-10
of Bobwhite quail, 45
of broiler breeder chickens, 32-34
of broiler chickens, 26-32
carbohydrates, 8
of ducks, 42-43
energy levels, 7-8
fatty acids, 13
of geese, 40-41
of Japanese quail, 44-45
of Leghorn-type breeders, 26
of Leghorn-type chickens, documentation of, 85-89
of Leghorn-type hens in egg production, 22-26
of Leghorn-type molting hens, 26
of Leghorn-type pullets, 19-22
minerals, 13-15
of ring-necked pheasants, 44
of turkeys, 35-39
vitamins, 15
O
Oats
amino acid digestibility coefficients, 74
fatty acid composition of, 75
metabolizable energy in, 113
nutrient composition of, 64-65, 67
Ochratoxin A, 79
Osteocalcin, 50
Oyster shells, 78
P
Pantothenic acid
broiler chicken requirements, 31
deficiency, 47, 48, 54
documentation of nutrient requirements, 87, 89, 95, 104,
105, 106, 108, 109, 111, 112
egg-type breeder requirements, 26
in energy metabolism, 54
in feeds, 63, 65
geese requirements, 41
maximum tolerance, 15
turkey requirements, 39
Paralysis, in nutrient deficiency, 31
Peanuts
amino acid digestibility coefficients, 74
fatty acid composition of, 75
nutrient composition of, 64-65, 67
Peas
amino acid composition of, 71, 72
nutrient composition of, 64-65, 67
Pelleted feeds, 15-16
for ducks, 42
thiamin in, 53
for turkeys, 35, 37
Perosis, 41, 53, 54, 55, 56
Phase feeding of Leghorn-type chickens, 24
Pheasants, 54
nutrient requirements for, 44, 109
Phenylalanine, 11
broiler chicken requirements, 29
deficiency, 46, 50
documentation of nutrient requirements, 92, 97, 101-102,
110
in feeds, 66-68
tyrosine interaction, 10
Phosphoric acid, element concentrations in, 78
Phosphorus
Bobwhite quail requirements, 45
broiler breeder chicken requirements, 33, 34
broiler chicken requirements, 29, 30
deficiency, 47, 56
digestibility, 14
documentation of nutrient requirements, 85, 88, 93-94, 97,
98, 103 , 105, 106, 107, 109, 111, 112
in egg production, 25
in feeds, 72, 64, 78
geese requirements, 41
in Leghorn-type pullet growth, 21
nutritional role of, 14
in phase-feeding program, 24
plant sources, 56
turkey requirements, 38
Phylloquinone, 15
source of, 52
Phytic acid, 38
Pigmentation
gossypol pigments in cottonseed oil, 69
mineral deficiency and, 57
nutrient deficiency in depigmentation, 46
pigments in feedstuffs, 17, 50
Potassium
broiler chicken requirements, 29, 30
deficiency, 47, 56-57
documentation of nutrient requirements, 86, 88, 94, 103
in feeds, 72, 64, 78
nutritional role of, 14
Poultry by-products
amino acid composition of, 67, 72
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INDEX
152
amino acid digestibility coefficients, 74
fatty acid composition of, 75
metabolizable energy in, 6, 114
nutrient composition of, 64-65
Proline
documentation of nutrient requirements, 93
Proline, broiler chicken requirements, 29
Protein
deficiencies, 46
documentation of nutrient requirements, 97, 98, 99, 105,
106, 107, 109, 110, 112
egg weight and, 24
in feeds, 72, 64, 66-68, 69
in measuring metabolizable energy, 5, 6
in phase-feeding program, 24
supplements, 69-70
See also Amino acids;
Protein requirements
Protein requirements
of broiler breeder chickens, 32-33, 34
of ducks, 42
of geese, 40
of Leghorn-type laying hens, 22-24, 88
of Leghorn-type pullets, 19-21, 85
of pheasants, 44
of quail, 45
of turkeys, 36, 37
Proximate analysis, 6-7
Pullets
broiler breeder, 32-33
Leghorn-type, 19-22
Pyridoxine
broiler chicken requirements, 32
deficiency, 47, 48, 54-55
documentation of nutrient requirements, 87, 89, 96, 104,
108, 111
in feeds, 63, 65
maximum tolerance, 15
in nervous system functioning, 54
Q
Quail, 54
Bobwhite, 45, 56, 112
Japanese, 44-45, 110-111
R
Rapeseed
metabolizable energy in, 113
as protein supplement, 69-70
Riboflavin
biochemical role of, 53
broiler chicken requirements, 30, 31
deficiency, 47, 48, 53
documentation of nutrient requirements, 87, 89, 95, 104,
105, 108, 109, 111, 112
egg-type breeder requirements, 26
in feeds, 63, 65
geese requirements, 41
maximum tolerance, 15
turkey requirements, 39
Rice, 113
amino acid composition of, 71
amino acid digestibility coefficients, 74
nutrient composition of, 64-65, 67
Rye, 9
nutrient composition of, 64-65, 67
S
Safety margin in requirements values, 1, 2
Safflower, nutrient composition of, 64-65, 67
Salt, 14
water intake and, 16
Selenium, 14, 50
broiler chicken requirements, 31
deficiency, 47, 48, 51, 52, 57
documentation of nutrient requirements, 86, 89, 94, 103,
107, 111
in feeds, 63, 65
sources of, 57
toxicity, 57, 58, 60
in vitamin E metabolism, 51
Serine
broiler chicken requirements, 29
documentation of nutrient requirements, 90, 110
in feeds, 66-68
glycine interaction, 10
Sesame
amino acid digestibility coefficients, 74
nutrient composition of, 64-65, 67-68
Silicon, 15
Silver, toxicity of, 60
Sodium
broiler breeder chicken requirements, 33
deficiency, 47, 57
documentation of nutrient requirements, 21, 86, 88, 94, 98,
103, 109
in egg production, 25
in feeds, 63, 65, 78
nutritional role of, 14
toxicity, 60
Sodium carbonate, element concentrations in, 78
Sodium chloride
documentation of nutrient requirements, 86, 107
to induce molt, 26
toxicity, 60
Sodium chlorine
documentation of nutrient requirements, 111
Sodium phosphate, element concentrations in, 78
Sodium sulfate, element concentrations in, 78
Sorghum
fatty acid composition, 75
metabolizable energy of, 6, 113
nutrient composition of, 64-65, 68-69
tannin content of, 6, 61, 68-69
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153
Soybeans/soybean meal
amino acid composition of, 71, 72
amino acid digestibility coefficients, 74
fatty acid composition of, 75
metabolizable energy in, 113
nutrient composition of, 64-65, 68
as protein supplement, 70
toxic elements in, 70
urease assays, 70
Standard reference diets, 80-81
Strontium, toxicity of, 60
Sulfaquinoxaline, 52
Sulfate
in broiler chicken diet, 28
toxicity of, 60
Sulfur amino acids, 63, 65, 78
in Bobwhite quail, 45
in broiler chickens, 28
in ducks, 43
in geese, 41
in quail, 45
in turkeys, 38
Sunflower
amino acid composition of, 71, 72
amino acid digestibility coefficients, 74
metabolizable energy in, 113
nutrient composition of, 64-65, 68
Supplementation
amino acids, 74
antibiotics, 18
antimicrobial additives, 18
of cereal grain diets, 9
choline, 55-56
fats, 6-7, 25
minerals, 14-15
protein, 69
selenium, 57
vitamin K, 52
vitamins, 15
of wheat diets, 69
Symptoms of nutrient deficiency
adrenal weight increase, 57
blood disorders, 49, 52, 53, 55, 57
bone abnormalities, 49, 51, 53, 54, 55, 56, 57
cervical paralysis, 55
dermatitis, 53, 54
diarrhea, 49
encephalomalacia, 51, 52
feather abnormalities, 44, 46, 49, 50, 51, 52, 53, 54, 55, 57
foot pad dermatitis, 50
gizzard ulcerations, 70
hyperthyroidism, 50
hypocalcemia, 51
immune system function, 50, 55
leg disorders in pheasants, 44
liver function, 46
muscle abnormalities, 46, 51, 57
nervous system dysfunction, 49, 50, 51, 52, 53, 54
pancreatic disorders, 57
pendulous crops, 54
pigmentation disorders, 46, 57
reproductive disorders, 50, 52, 54
rickets, 56
secretory membrane dysfunction, 50
skin lesions, 49, 50, 54
thyroid disorders, 57
tongue deformity, 50
T
Temperature, environmental
feed intake and, 8, 9, 22
phosphorus in tolerance to, 25
in requirements data, 2
turkey maintenance energy and, 37
water intake and, 16
Thiamin
broiler chicken requirements, 31-32
deficiency, 47, 48, 52-53
documentation of nutrient requirements, 87, 89, 96, 104, 111
in feeds, 63, 65
maximum tolerance, 15
Threonine, 11
broiler chicken requirements, 29
documentation of nutrient requirements, 85, 88, 92, 97,
102, 110
in feeds, 66-68, 71-73
D/DL-α-Tocopheryl/Tocopheryl acetate, 15
Toxicity
amino acid, 11
calcium, 44
definition, 58
gossypol pigments in cottonseed oil, 69
of inorganic elements, 57, 58-60
methionine, 11
mycotoxin, 78-79
vitamin A, 50
vitamin D, 51
vitamin E, 52
Tricothecenes, 79
Triticale, 113
amino acid composition of, 71
nutrient composition of, 64-65, 68
Tryptophan
broiler breeder requirements
broiler chicken requirements, 29
deficiency, 46
documentation of nutrient requirements, 92-93, 102, 107,
110
duck requirements, 43
egg weight and, 25
in feeds, 66-68, 71-73
as niacin source, 11, 53
Tungsten, toxicity of, 60
Turkeys
amino acid requirements, 36, 37-38
breeding diets, 38, 39
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INDEX
154
documentation of nutrient requirements, 99-105
egg production, 37, 38
energy consumption, 37
magnesium requirements, 38
manganese requirements, 38
mineral requirements, 38
nutrient requirements, 35-36
nutrient-to-energy ratios in feeds, 8
nutritional modeling for, 1
pelleted feeds for, 35, 37
phosphorus requirements, 38
protein requirements, 36, 37
vitamin requirements, 38-39
water intake, 16
zinc requirements, 38
See also Symptoms of nutrient deficiency
Tyrosine, 11
broiler chicken requirements, 29
deficiency, 46
documentation of nutrient requirements, 92, 101, 110
in feeds, 66-68
phenylalanine interaction, 10
U
Urinary energy, 5
V
Valine, 11
broiler chicken requirements, 29
deficiency, 46
documentation of nutrient requirements, 85, 88, 93, 97,
102, 107, 110
duck requirements, 43
in feeds, 66-68
Vanadium, toxicity of, 58, 60
Vegetable oils, 12, 13
Vitamin A, 15
broiler chicken requirements, 31
deficiency, 47, 48, 50
documentation of nutrient requirements, 86, 89, 94, 103,
105, 111, 112
maximum tolerance, 15
toxicity, 50
turkey requirements, 38
Vitamin B12, 15
broiler chicken requirements, 31
deficiency, 47, 48, 55
documentation of nutrient requirements, 87, 89, 95, 104
egg-type breeder requirements, 26
in feeds, 63, 65
role of, 55
Vitamin C
bird stress and, 15
maximum tolerance, 15
Vitamin D, 15
broiler chicken requirements, 30, 31
in calcium metabolism, 50, 51
deficiency, 47, 48, 50-51, 51
documentation of nutrient requirements, 86, 89, 94-95,
103, 105, 107, 109, 111
maximum tolerance, 15
metabolism, 50
nutritional role of, 50
sources of, 50
toxicity, 51
turkey requirements, 38-39
Vitamin E, 15
broiler chicken requirements, 30, 31
deficiency, 47, 48, 51-52
documentation of nutrient requirements, 95, 103, 105, 108
documentation of nutrient requirements, 89, 87
in feeds, 63, 65
maximum tolerance, 15
metabolism, 51
toxicity, 52
turkey requirements, 39
in vitamin A excess, 50
Vitamin K, 15
broiler chicken requirements, 31
deficiency, 47, 48, 52
documentation of nutrient requirements, 87, 89, 95, 103, 108
maximum tolerance, 15
nutritional role of, 52
sources of, 52
turkey requirements, 39
Vitamins, 15
amino acids in synthesis of, 11
antagonistic interactions, 50
broiler breeder chicken requirements, 33
broiler chicken requirements, 27, 30, 31-32
deficiencies, 47, 48, 50-56
documentation of nutrient requirements, 86-87, 89, 94-96,
103-104, 105, 107-108, 111
duck requirements, 43
geese requirements, 41
Leghorn-type chicken requirements, 20, 21, 22, 25, 26
maximum tolerances, 15
pheasant requirements, 44
as supplements, 15
turkey requirements, 38-39
W
Water
delivery systems, 16
deprivation of, 16
intake determinants, 15-16
intoxication, 16
medication in, 16
requirements, 15, 16
total dissolved solids in, 17
trace minerals in, 16-17
Wheat
amino acid composition of, 71
amino acid digestibility coefficients, 74
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INDEX
155
fatty acid composition of, 75
metabolizable energy of, 69, 113
nutrient composition of, 64-65, 68
Whey, nutrient composition of, 64-65, 68
X
Xanthophylls, 17
role of, 3
Y
Yeasts, nutrient composition of, 64-65, 68
Zearalenone, 79
Zeaxanthin, 17
Zinc, 14
broiler chicken requirements, 29, 30
deficiency, 47, 48, 57
documentation of nutrient requirements, 86, 88, 94, 103,
107, 109, 111
in feeds, 63, 65, 78
to induce molt, 26
pheasant requirements, 44
toxicity, 60
turkey requirements, 38
Z
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INDEX
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Nutrient Requirements of Poultry: Ninth Revised Edition, 1994
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RELATED PUBLICATIONS
157
Other Titles in the Series
Nutrient Requirements of Poultry, 9th Rev. Ed., 1994
Nutrient Requirements of Fish, 1993
Nutrient Requirements of Horses, 5th Rev. Ed., 1989
Nutrient Requirements of Dairy Cattle, 6th Rev. Ed., Update 1989
Nutrient Requirements of Swine, 9th Rev. Ed., 1988
Nutrient Requirements of Cats, Rev. Ed., 1986
Nutrient Requirements of Sheep, 6th Rev. Ed., 1985
Nutrient Requirements of Dogs, Rev. Ed., 1985
Nutrient Requirements of Beef Cattle, 6th Rev. Ed., 1984
Nutrient Requirements of Warm Water Fishes and Shellfishes, Rev. Ed., 1983
Nutrient Requirements of Mink and Foxes, Rev. Ed., 1982
Nutrient Requirements of Coldwater Fishes, 1981
Nutrient Requirements of Goats, 1981
Nutrient Requirements of Nonhuman Primates, 1978
Nutrient Requirements of Rabbits, 2nd Rev. Ed., 1977
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