UseAlageAquaticFeed2009 FAO

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FAO

ISSN 2070-7010
FISHERIES AND
AQUACULTURE

531
TECHNICAL
PAPER

531
Use of algae and aquatic
macrophytes as feed in
small-scale aquaculture
A review

Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review


While the contribution of small-scale aquaculture (SSA) to rural development is
generally recognized, until now there has been no systematic assessment to clearly
measures its contribution. The FAO Expert Workshop on Methods and Indicators for
Evaluating the Contribution of Small-scale Aquaculture to Sustainable Rural Development
held in Nha Trang, Viet Nam, from 24 to 28 November 2009, attempted to develop
an indicator system to measure the contribution of SSA. The workshop used a
number of processes and steps in the developping the indicator system, including:
(i) understanding the subject of measurements; (ii) identifying an analytical framework
and ratting criteria (iii) developing a list of SSA contributions; (iv) categorizing the contributions;
(v) devising and organizing the indicators of contribution; and (vi) measuring the indicators.
The major outcome was the development, through an iterative process, of an indicator
system which can provide a good measure of the contribution of SSA based on agreed
criteria (accuracy, measurability and efficiency) and the sustainable livelihood
approach analytical framework which consists of five capital assets (human, financial,
physical, social and natural) and can be used for various livelihoods options.

FAO
Cover photographs:
Left: Woman collecting water chestnut fruits from a floodplain, Rangpur, Bangladesh (courtesy of
Mohammad R. Hasan).
Right top to bottom: Sale of water spinach leaves, Ho Chi Minh City, Viet Nam (courtesy of William
Leschen). Woman carrying water spinach leaves after harvest, Beung Cheung Ek wastewater lake,
Phnom Penh, Cambodia (courtesy of William Leschen). Back side of a lotus leave, photograph taken in a
floodplain, Rangpur, Bangladesh (courtesy of Mohammad R. Hasan).
Use of algae and aquatic
FAO
FISHERIES AND
AQUACULTURE

macrophytes as feed in
TECHNICAL
PAPER

531
small-scale aquaculture
A review

by

Mohammad R. Hasan
Aquaculture Management and Conservation Service
Fisheries and Aquaculture Management Division
FAO Fisheries and Aquaculture Department
Rome, Italy

and

Rina Chakrabarti
University of Delhi
Delhi, India

FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS


Rome, 2009
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necessarily reflect the views of FAO.

ISBN 978-92-5-106420-7

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© FAO 2009
iii

Preparation of this document

Recognizing the increasing importance of the use of aquatic macrophytes as feed in


small-scale aquaculture, the global review on this topic was undertaken as a part of the
regular work programme of the Fisheries and Aquaculture Department of the Food and
Agriculture Organization of the United Nations (FAO) by the Aquaculture Management
and Conservation Service ‘Study and analysis of feed and nutrients (including fertilizers)
for sustainable aquaculture development’. This was carried out under the programme entity
‘Monitoring, Management and Conservation of Resources for Aquaculture Development’.
The manuscript was edited for technical content by Michael B. New. For consistency and
conformity, scientific and English common names of fish species were used from FishBase
(www.fishbase.org/home.htm). Most of the photographs in the manuscripts were provided
by the first author. Where this is not the case, due acknowledgements are made to the
contributor(s) or the source(s).
Special thanks are due to Dr Albert G.J. Tacon (Universidad de Las Palmas de Gran
Canaria, Spain), Dr M.A.B. Habib (Bangladesh Agricultural University, Bangladesh), Md.
Ghulam Kibria (Ministry of Fisheries and Marine Resources, Namibia) and Dr Khondker
Moniruzzaman (University of Dhaka, Bangladesh) who kindly provided papers and
information. The Royal Netherlands Embassy in Dhaka, Bangladesh is acknowledged for
kindly providing the reports of the Duckweed Research Project.
We acknowledge Ms Tina Farmer and Ms Françoise Schatto for their assistance in quality
control and FAO house style and Mr Juan Carlos Trabucco for layout design. The publishing
and distribution of the document were undertaken by FAO, Rome.
Mr Jiansan Jia, Service Chief, and Dr Rohana P. Subasinghe, Senior Fishery Resources
Officer (Aquaculture), Aquaculture Management and Conservation Service of the FAO
Fisheries and Aquaculture Department are also gratefully acknowledged for their support.
iv

Abstract

This technical paper presents a global review on the use of aquatic macrophytes as feed
for farmed fish, with particular reference to their current and potential use by small-scale
farmers. The review is organized under four major divisions of aquatic macrophytes: algae,
floating macrophytes, submerged macrophytes and emergent macrophytes. Under floating
macrophytes, Azolla, duckweeds and water hyacinths are discussed separately; the remaining
floating macrophytes are grouped together and are reviewed as ‘other floating macrophytes’.
The review covers aspects concerned with the production and/or cultivation techniques and
use of the macrophytes in their fresh and/or processed state as feed for farmed fish. Efficiency
of feeding is evaluated by presenting data on growth, food conversion and digestibility
of target fish species. Results of laboratory and field trials and on-farm utilization of
macrophytes by farmed fish species are presented. The paper provides information on the
different processing methods employed (including composting and fermentation) and results
obtained to date with different species throughout the world with particular reference to
Asia. Finally, it gives information on the proximate and chemical composition of most
commonly occurring macrophytes, their classification and their geographical distribution
and environmental requirements.

Hasan, M.R.; Chakrabarti, R.


Use of algae and aquatic macrophytes as feed in small-scale aquaculture: a review.
FAO Fisheries and Aquaculture Technical Paper. No. 531. Rome, FAO. 2009. 123p.


Contents

Preparation of this document iii


Abstract iv
Abbreviations and acronyms vii

Introduction 1

1. Algae 3
Classification 3
Characteristics 4
Production 7
Chemical composition 8
Use as aquafeed 8

2. Floating aquatic macrophytes – Azolla 17


Classification 17
Characteristics 17
Production 19
Chemical composition 21
Use as aquafeed 21

3. Floating aquatic macrophytes – duckweeds 29


Classification 29
Characteristics 31
Production 34
Chemical composition 40
Use as aquafeed 43

4. Floating aquatic macrophytes – water hyacinths 53


Classification 53
Characteristics 53
Production 54
Chemical composition 55
Use as aquafeed 55

5. Floating aquatic macrophytes – others 67


Classification 67
Characteristics 67
Production 68
Chemical composition 68
Use as aquafeed 70
vi

6. Submerged aquatic macrophytes 75


Classification 75
Characteristics 75
Production 76
Chemical composition 76
Use as aquafeed 78

7. Emergent aquatic macrophytes 89


Classification 89
Characteristics 90
Production 91
Chemical composition 91
Use as aquafeed 93

8. Conclusions 95
Algae 95
Azolla 95
Duckweeds 96
Water hyacinths 96
Other floating macrophytes 97
Submerged macrophytes 98
Emergent macrophytes 99

References 101
Annex 1 Essential amino acid composition of aquatic macrophytes 119
Annex 2 Periphyton 123
vii

Abbreviations and acronyms

APD Apparent Protein Digestibility


BFRI Bangladesh Fishery Research Institute
BW Body Weight
DM Dry Matter basis
DW Dry weight
DWRP Duckweed Research Project (Bangladesh)
EAA Essential Amino Acid
FCR Feed Conversion Ratio
FW Fresh Weight
MAEP Mymensingh Aquaculture Extension Project
MP Muriate of Potash
NFE Nitrogen-Free Extract
NGO Non-governmental organization
PRISM Project in Agriculture, Rural Industry Science and Medicine (an
NGO)
SGR Specific Growth Rate
TKN Total Kjeldahl Nitrogen
TSP Triple Super Phosphate
UASB Upflow Anaerobic Sludge Blanket Reactor
2,4-D 2,4-Dichhlorophenoxyacetic acid


Introduction

Using feeds in aquaculture (sometimes referred to as aquafeeds) generally increases


productivity. However, to maximize cost-effectiveness, it is particularly useful in
small-scale aquaculture to utilize locally available materials, either as ingredients (raw
materials) in compound aquafeeds or as sole feedstuffs.
There is also a vital need to seek effective ingredients that can either partially or
totally replace marine ingredients as protein sources in animal feedstuffs generally, in
particular in aquafeeds. While this broad topic is not dealt with in this review, many
introductions to the literature of the past two decades are available, including New and
Csavas (1995), Tacon (1994; 2004;), Tacon, Hasan and Subasinghe (2006), Tacon and
Metain (2008), New and Wijkstrom (2002), FAO (2008) and Huntington and Hasan
(2009).
This review deals with the characteristics of aquatic raw materials for use as feeds in
small-scale aquaculture, namely algae (principally macro-algae – commonly referred to
as seaweeds) and aquatic macrophytes. Aquatic macrophytes are aquatic plants that are
large enough to be seen by the naked eye. They grow in or near water and are floating,
submerged, or emergent.
Information includes current and potential usage of these materials by small-scale
aquafarmers for target fish and crustaceans, together with details on their classification,
characteristics (including such factors as their natural distribution and environmental
requirements), production and chemical composition.
The review has been divided into seven major sections: one dealing with algae;
four sections on floating macrophytes (namely Azolla, duckweeds, water hyacinths
and others); a section on submerged macrophytes; and one on emergent macrophytes.
Finally, the review contains a concluding section which summarizes previous
chapters.


1. Algae

Algae have been used in animal and human diets since very early times. Filamentous
algae are usually considered as ‘macrophytes’ since they often form floating masses that
can be easily harvested, although many consist of microscopic, individual filaments
of algal cells. Algae also form a component of periphyton, which not only provides
natural food for fish and other aquatic animals but is actively promoted by fishers and
aquaculturists as a means of increasing productivity. This topic is not dealt with in
this section, since periphyton is not solely comprised of algae and certainly cannot be
regarded as macroalgae. However, some ancillary information on this topic is provided
in Annex 2 to assist with further reading. Marine ‘seaweeds’ are macro-algae that have
defined and characteristic structures.
Microalgal biotechnology only really began to develop in the middle of the last
century but it has numerous commercial applications. Algal products can be used
to enhance the nutritional value of food and animal feed owing to their chemical
composition; they play a crucial role in aquaculture. Macroscopic marine algae
(seaweeds) for human consumption, especially nori (Porphyra spp.), wakame (Undaria
pinnatifida), and kombu (Laminaria japonica), are widely cultivated algal crops. The
most widespread application of microalgal culture has been in artificial food chains
supporting the husbandry of marine animals, including finfish, crustaceans, and
molluscs.
The culture of seaweed is a growing worldwide industry, producing 14.5 million
tonnes (wet weight) worth US$7.54 billion in 2007 (FAO, 2009). The use of aquatic
macrophytes in treating sewage effluents has also shown potential. In recent years,
macroalgae have been increasingly used as animal fodder supplements and for the
production of alginate, which is used as a binder in feeds for farm animals. Laboratory
investigations have also been carried out to evaluate both algae and macroalgae as
possible alternative protein sources for farmed fish because of their high protein content
and productivity.
Microalgae and macroalgae are also used as components in polyculture systems
and in remediation; although these topics are not covered in this paper, information
on bioremediation is contained in many publications, including Msuya and Neori
(2002), Zhou et al. (2006) and Marinho-Soriano (2007). Red seaweed (Gracilaria spp.)
and green seaweed (Ulva spp.) have been found to suitable species for bioremediation.
The use of algae in integrated aquaculture has also been recently reviewed by Turan
(2009).

1.1 Classification
The classification of algae is complex and somewhat controversial, especially concerning
the blue-green algae (Cyanobacteria), which are sometimes known as blue-green
bacteria or Cyanophyta and sometimes included in the Chlorophyta. These topics are
not covered in detail this document. However, the following provides a taxonomical
outline of algae.
Archaeplastida
• Chlorophyta (green algae)
• Rhodophyta (red algae)
• Glaucophyta
Rhizaria, Excavata
• Chlorarachniophytes
 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

• Euglenids
Chromista, Alveolata
• Heterokonts
• Bacillariophyceae (diatoms)
• Axodine
• Bolidomonas
• Eustigmatophyceae
• Phaeophyceae (brown algae)
• Chrysophyceae (golden algae)
• Raphidophyceae
• Synurophyceae
• Xanthophyceae (yellow-green algae)
• Cryptophyta
• Dinoflagellates
• Haptophyta

The following sections discuss the characteristics and use of both ‘true’ algae and the
Cyanophyta, hereinafter referred to as ‘blue-green algae’).

1.2 Characteristics
Filamentous algae and seaweeds have an extremely wide panorama of environmental
requirements, which vary according to species and location. Ecologically, algae are
the most widespread of the photosynthetic plants, constituting the bulk of carbon
assimilation through microscopic cells in marine and freshwater.
The environmental requirements of algae are not discussed in detail in this document.
However, the most important parameters regulating algal growth are nutrient quantity
and quality, light, pH, turbulence, salinity and temperature. Macronutrients (nitrate,
phosphate and silicate) and micronutrients (various trace metals and the vitamins
thiamine (B1), cyanocobalamin (B12) and biotin) are required for algal growth (Reddy
et al., 2005). Light intensity plays an important role, but the requirements greatly
vary with the depth and density of the algal culture. The pH range for most cultured
algal species is between 7 and 9; the optimum range is 8.2–8.7. Marine phytoplankton
are extremely tolerant to changes in salinity. In artificial culture, most grow best at
a salinity that is lower than that of their native habitat. Salinities of 20–24 ppt are
found to be optimal. Lapointe and Connell (1989) suggested that the growth of the
green filamentous alga Cladophora was limited by both nitrogen and phosphorus, but
the former was the primary factor. Hall and Payne (1997) found that another green
filamentous alga, Hydrodictyon reticulatum, had a relatively low requirement for
dissolved inorganic nitrogen in comparison with other species. Rafiqul, Jalal and Alam
(2005) found that the optimum environment for Spirulina platensis under laboratory
conditions was 32 ºC, 2 500 lux and pH 9.0. Further information on the environmental
requirements of algae cultured for use in aquaculture hatcheries is contained in Lavens
and Sorgeloos (1996). The environmental requirements of cultured seaweeds are
discussed by McHugh (2002, 2003).
A brief description of some of the filamentous algae and seaweeds that have been used
for feeding fish, as listed in Tables 1.1–1.3, is provided in the following subsections.

1.2.1 Filamentous algae


Filamentous algae are commonly referred to as ‘pond scum’ or ‘pond moss’ and
form greenish mats upon the water surface. These stringy, fast-growing algae can
cover a pond with slimy, lime-green clumps or mats in a short period of time, usually
beginning their growth along the edges or bottom of the pond and ‘mushrooming’ to
the surface. Individual filaments are a series of cells joined end to end which give the
Algae 

thread-like appearance. They also form fur-like growths on submerged logs, rocks and
even on animals. Some forms of filamentous algae are commonly referred to as ‘frog
spittle’ or ‘water net’.
Spirulina, which is a genus of cyanobacteria that is also considered to be a
filamentous blue-green algae, is cultivated around the world and used as a human
dietary supplement, as well as a whole food. It is also used as a feed supplement in the
aquaculture, aquarium, and poultry industries (Figure 1.1).

Figure 1.1 Figure 1.2


Spirulina sp. Spirogyra sp.

Source: scienceblogs.com/energy/2008/08/big_
Source: Wim van Egmont©
energy_fr...

Spirogyra, one of the commonest green filamentous algae (Figure 1.2), is named
because of the helical or spiral arrangement of the chloroplasts. There are more than
400 species of Spirogyra in the world. This genus is photosynthetic, with long bright
grass-green filaments having spiral-shaped chloroplasts. It is bright green in the spring,
when it is most abundant, but deteriorates to yellow. In nature, Spirogyra grows in
running streams of cool freshwater, and secretes a coating of mucous that makes it
feel slippery. This freshwater alga is found in shallow ponds, ditches and amongst
vegetation at the edges of large lakes. Under favourable conditions, Spirogyra forms
dense mats that float on or just beneath the surface of the water. Blooms cause a grassy
odour and clog filters, especially at water treatment facilities.
Cladophora (Figure 1.3) is a green filamentous algae that is a member of the
Ulvophyceae and is thus related to the sea lettuce (Ulva spp.). The genus Cladophora
has one of the largest number of species within the macroscopic green algae and is
also among the most difficult to classify taxonomically. This is mainly due to the
great variations in appearance, which are significantly affected by habitat, age and
environmental conditions. These algae, unlike Spirogyra, do not conjugate (form
bridges between cells) but simply branch.

Figure 1.3 Figure 1.4


Cladophora sp. Water net (Hydrodictyon sp.)

Source: Biopix.dk© Source: silicasecchidisk.conncoll.edu


 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

Another green filamentous alga, Hydrodictyon, commonly known as ‘water net’,


belongs to the family Hydrodictyaceae and prefers clean, eutrophic water. Its name
refers to its shape, which looks like a netlike hollow sack (Figure 1.4) and can grow
up to several decimetres.

1.2.2 Seaweeds
Ulva are thin flat green algae growing from a discoid holdfast that may reach 18 cm or
more in length, though generally much less, and up to 30 cm across. The membrane is
two cells thick, soft and translucent and grows attached (without a stipe) to rocks by
a small disc-shaped holdfast. The water lettuce (Ulva lactuca) is green to dark green
in colour (Figure 1.5). There are other species of Ulva that are similar and difficult to
differentiate.

Figure 1.5 Figure 1.6


Sea lettuce (Ulva lactuca) Eucheuma cottonii

Source: Mandy Lindeberg (www.seaweedsofalaska.com) Source: www.actsinc.biz/seaweed.html

It is important to recognize that the genera Eucheuma and Kappaphycus are


normally grouped together; their taxonomical classification is contentious. These are
red seaweeds and are often very large macroalgae that grow rapidly. The systematics
and taxonomy of Kappaphycus and Eucheuma (Figure 1.6) is confused and difficult, due
to morphological plasticity, lack of adequate characters to identify species and the use
of commercial names of convenience. These taxa are geographically widely dispersed
through cultivation (Zuccarello et al., 2006). These red seaweeds are widely cultivated,
particularly to provide a source of carageenan, which is used in the manufacture of
food, both for humans and other animals.
Gracilaria is another genus of red algae (Figure 1.7), most well-known for its
economic importance as a source of agar, as well as its use as a food for humans.

Figure 1.7 Figure 1.8


Gracilaria sp. Porphyra tenera

Source: Eric Moody© (Wikipedia) Source: http://www.fao.org/fishery/species/search/en


Algae 

The red seaweed Porphyra (Figure 1.8) is known by many local names, such as laver
or nori, and there are about 100 species. This genus has been cultivated extensively in
many Asian countries and is used to wrap the rice and fish that compose the Japanese
food sushi, and the Korean food gimbap. It is also used to make the traditional Welsh
delicacy, laverbread.

1.3 Production
As in the case of their environmental conditions, the methods for culturing filamentous
algae and seaweeds vary widely, according to species and location. This topic is not
covered in this review but there are many publications available on algal culture
generally, such as the FAO manual on the production of live food for aquaculture by
Lavens and Sorgeloos (1996). Concerning seaweed culture, the following summary
of the techniques used has been has been extracted from another FAO publication
(McHugh, 2003) and further reading on seaweed culture can also be found in McHugh
(2002).

Some seaweeds can be cultivated vegetatively, others only by going through a separate
reproductive cycle, involving alternation of generations.
In vegetative cultivation, small pieces of seaweed are taken and placed in an
environment that will sustain their growth. When they have grown to a suitable size they
are harvested, either by removing the entire plant or by removing most of it but leaving
a small piece that will grow again. When the whole plant is removed, small pieces are cut
from it and used as seedstock for further cultivation. The suitable environment varies
among species, but must meet requirements for salinity of the water, nutrients, water
movement, water temperature and light. The seaweed can be held in this environment
in several ways: pieces of seaweed may be tied to long ropes suspended in the water
between wooden stakes, or tied to ropes on a floating wooden framework (a raft);
sometimes netting is used instead of ropes; in some cases the seaweed is simply placed
on the bottom of a pond and not fixed in any way; in more open waters, one kind of
seaweed is either forced into the soft sediment on the sea bottom with a fork-like tool,
or held in place on a sandy bottom by attaching it to sand-filled plastic tubes.
Cultivation involving a reproductive cycle, with alternation of generations, is
necessary for many seaweeds; for these, new plants cannot be grown by taking
cuttings from mature ones. This is typical for many of the brown seaweeds, and
Laminaria species are a good example; their life cycle involves alternation between a
large sporophyte and a microscopic gametophyte-two generations with quite different
forms. The sporophyte is what is harvested as seaweed, and to grow a new sporophyte
it is necessary to go through a sexual phase involving the gametophytes. The mature
sporophyte releases spores that germinate and grow into microscopic gametophytes.
The gametophytes become fertile, release sperm and eggs that join to form embryonic
sporophytes. These slowly develop into the large sporophytes that we harvest. The
principal difficulties in this kind of cultivation lie in the management of the transitions
from spore to gametophyte to embryonic sporophyte; these transitions are usually
carried out in land-based facilities with careful control of water temperature, nutrients
and light. The high costs involved in this can be absorbed if the seaweed is sold as
food, but the cost is normally too high for production of raw material for alginate
production.
Where cultivation is used to produce seaweeds for the hydrocolloid industry (agar
and carrageenan), the vegetative method is mostly used, while the principal seaweeds
used as food must be taken through the alternation of generations for their cultivation.
 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

1.4 Chemical composition


A summary of the chemical composition of various filamentous algae and seaweeds is
presented in Table 1.1. Algae are receiving increasing attention as possible alternative
protein sources for farmed fish, particularly in tropical developing countries, because
of their high protein content (especially the filamentous blue-green algae).
The dry matter basis (DM) analyses reviewed in Table 1.1 show that the protein
levels of filamentous blue green algae ranged from 60–74 percent. Those for filamentous
green algae were much lower (16–32 percent). The protein contents of green and red
seaweeds were quite variable, ranging from 6–26 percent and 3–29 percent respectively.
The levels reported for Eucheuma/ Kappaphycus were very low, ranging from 3–10
percent, but the results for Gracilaria, with one exception, were much higher (16–20
percent). The one analysis for Porphyra indicated that it had a protein level (29 percent)
comparable to filamentous green algae. Some information on the amino acid content of
various aquatic macrophytes is contained in Annex 1.
The lipid levels reported for Spirulina (Table 1.1), with one exception (Olvera-
Novoa et al. (1998), were between and 4 and 7 percent. Those for filamentous green
algae varied more widely (2–7 percent). The lipid contents of both green (0.3–3.2
percent) and red seaweeds (0.1–1.8 percent) were generally much lower than those of
filamentous algae. The ash content of filamentous blue-green algae ranged from 3–11
percent but those of filamentous green algae were generally much higher, ranging from
just under 12 percent to one sample of Cladophora that had over 44 percent. The ash
contents of green seaweeds ranged from 12–31 percent. Red seaweeds had an extremely
wide range of ash contents (4 to nearly 47 percent) and generally had higher levels than
the other algae shown in Table 1.1.

1.5 Use as aquafeed


Several feeding trials have been carried out to evaluate algae as fish feed. Algae have
been used fresh as a whole diet and dried algal meal has been used as a partial or
complete replacement of fishmeal protein in pelleted diets.

1.5.1 Algae as major dietary ingredients


A summary of the results of selected growth studies on the use of fresh algae or dry
algae meals as major dietary ingredients for various fish species and one marine shrimp
is presented in Table 1.2. Dietary inclusion levels in these studies varied from 5 to 100
percent. Fishmeal-based dry pellets or moist diets were used as control diets.
The results of the earlier growth studies showed that the performances of fish fed
diets containing 10–20 percent algae or seaweed meal were similar to those fed fishmeal
based standard control diet. The responses were apparently similar for most of the
fish species tested. These inclusion levels effectively supplied only about 3–5 percent
protein of the control diet. In most cases, these control diets contained about 26–47
percent crude protein. This shows that only about 10–15 percent of dietary protein
requirement can be met by algae without compromising growth and food utilization.
There was a progressive decrease in fish performance when dietary incorporation of
algal meal rose above 15–20 percent. However, although reduced growth responses
were recorded with increasing inclusion of algae in the diet, the results of feeding trials
with filamentous green algae for O. niloticus and T. zillii indicated that SGR of 60–80
percent of the control diet could be achieved with dietary inclusion levels as high as
50–70 percent.
Recent work by Kalla et al. (2008) appears to indicate that the addition of Porphyra
spheroplasts to a semi-purified red seabream diet improved SGR. In addition, Valente
et al. (2006) recorded improvements in SGR when dried Gracilaria busra-pastonis
replaced 5 or 10 percent of a fish protein hydrolysate diet for European seabass.
Table 1.1
Chemical analyses of some common algae and seaweeds
Algae

Algae/ seaweed Moisture Proximate composition1 Minerals1 Reference


(percent) (percent DM) (percent DM)
CP EE Ash CF NFE Ca P
Filamentous blue-green algae
Spirulina maxima, spray dried powder 6.0 63.8 5.3 9.6 n.s. n.s. n.s. Henson (1990)
Spirulina, commercial dry powder 3-6 60.0 5.0 7.0 7.0 21.0 n.s. n.s. Habib et al. (2008)
Spirulina spp., dry powder n.s. 55-70 4-7 3-11 3-7 n.s. n.s. Habib et al. (2008)
Spirulina maxima, dry powder, Mexico 10.2 73.7 0.7 10.5 2.1 13.0 n.s. n.s. Olvera-Nova et al. (1998)
Filamentous green algae
Spirogyra spp., fresh, USA 95.2 17.1 1.8 11.7 10.02 n.s. n.s. Boyd (1968)
Cladophora glomerata, meal, Scotland 1.6 31.6 5.2 23.6 11.2 28.4 n.s. n.s. Appler and Jauncey (1983)
Cladophora sp., fresh, USA3 90.5 15.8 2.1 44.3 13.3 24.54 n.s. n.s. Pine, Anderson and Hung (1989)
2
Hydrodictyon reticulatum, fresh, USA 96.1 22.8 7.1 11.9 18.1 n.s. n.s. Boyd (1968)
Hydrodictyon reticulatum, meal, Belgium 5.7 27.7 1.9 32.6 14.9 22.9 n.s. n.s. Appler (1985)
Green seaweeds
Ulva reticulata, fresh, Tanzania n.s. 25.7 n.s. 18.3 38.5 n.s. 0.1 Msuya and Neori (2002)
Ulvaria oxysperma, dried, Brazil 16-20 6-10 0.5-3.2 17-31 3-12 n.s. n.s. Pádua, Fontoura and Mathias (2004)
Ulva lactuca, dried, Brazil 15-18 15-18 1.2-1.8 12-13 9-12 n.s. n.s. Pádua, Fontoura and Mathias (2004)
Ulva fascita, dried, Brazil 18-20 13-16 0.3-1.9 17-20 9-11 n.s. n.s. Pádua, Fontoura and Mathias (2004)
Red seaweeds
Eucheuma cottonii, fresh, Indonesia 91.3 4.9 0.4 43.5 8.4 42.84 0.5 0.2 Tacon et al. (1990)
Eucheuma cottonii, dry powder, Malaysia 10.6 9.8 1.1 46.2 5.9 37.04 0.3 n.s. Matanjun et al. (2009)
Eucheuma denticulatum, fresh, Tanzania n.s. 7.6 n.s. 46.6 22.3 n.s. <0.1 Msuya and Neori (2002)
Kappaphycus alvarezü, oven dried meal, Philippines 10.1 3.2 0.6 18.1 5.9 72.24 n.s. n.s. Peñaflorida and Golez (1996)
Gracilaria heteroclada, oven dried meal, Philippines 9.3 17.3 1.8 21.7 4.6 54.6 n.s. n.s. Peñaflorida and Golez (1996)
Gracilaria lichenoides, fresh, Indonesia 88.1 15.6 1.2 36.7 6.6 39.94 0.8 0.3 Tacon et al. (1990)
Gracilaria sp., sun-dried meal, Thailand 7.2 19.9 0.1 31.4 4.9 43.7 n.s. n.s. Briggs and Funge-Smith (1996)
Gracilaria crassa, fresh, Tanzania 13.2 n.s. 15.0 38.7 n.s. <0.1 Msuya and Neori (2002)
Porphyra purpurea, meal, England 4.7 28.7 0.4 4.1 6.7 60.14 n.s. n.s. Davies, Brown and Camilleri (1997)
1
DM = dry matter; CP = crude protein; EE = ether extract; CF = crude fibre; NFE = nitrogen free extract; Ca = calcium; P = phosphorus
2
Cellulose
3
Mean of proximate composition values of algae collected from flowing and static water
4
Adjusted or calculated; not as cited in original publication

10

Table 1.2
Performance of various fish species fed fresh algae or dried algal meal
Algae/ fish species Rearing Rearing Control diet Composition of test diet Inclusion Fish size SGR SGR as FCR References
system days level (g) (percent) percent of
(percent) control
Filamentous green algae
Cladophora glomerata/ Nile Laboratory 56 Fish meal based 5, 10, 15 and 20 percent 16.1 1.88-2.09 3.11 97.5 1.21 Appler and
tilapia (Oreochromis niloticus) recirculatory pellet (30 protein of control feed Jauncey
system percent protein) replaced by algal meal and (1983)
one diet prepared by algal 32.3 2.80 87.8 1.42
meal as the only sources
of protein (25 percent 48.4 2.77 86.8 1.51
protein) 64.5 2.06 64.6 2.09
82.5 1.85 58.0 2.33
Hydrodictyon reticulatum/ Laboratory 50 Fish meal based 5, 10, 15 and 20 percent 19.2 0.92-1.04 2.22 91.7 1.83 Appler
Nile tilapia (Oreochromis recirculatory pellet (30 protein of control feed 38.3 1.85 76.4 2.18 (1985)
niloticus) system percent protein) replaced by algal meal and
one diet prepared by algal 57.5 1.48 61.2 2.49
meal as the only sources 70.6 1.52 62.8 2.63
of protein (25 percent 98.5 1.07 44.2 3.60
protein)
Hydrodictyon reticulatum/ Laboratory 50 Fishmeal based 5, 10, 15 and 20 percent 19.2 0.91-1.16 2.04 107.9 2.09 Appler
Redbelly tilapia (Tilapia zillii) recirculatory pellet (30 protein of control feed 38.3 1.73 91.5 (1985)
system percent protein) replaced by algal meal and
one diet prepared by algal 57.5 1.45 76.7
meal as the only sources 70.6 1.44 76.2
of protein (25 percent 98.5 1.05 55.6
protein)
Filamentous blue-green algae
Spirulina/ Java tilapia Indoor static 25 Fishmeal based 11 percent fishmeal 11.0 7.4-8.3 1.96 101.0 - Chow and
(Oreochromis mossambicus) tank moist diet (26 replaced by Spirulina meal Woo (1996)
percent protein)
Seaweeds
Porphyra purpurea/ thick- Flow-through 70 Fishmeal based 4.5 and 9.0 percent protein 16.5 1.15 2.65 88.6 2.06 Davies,
lipped grey mullet (Chelon system pellet (47 of control feed replaced by 33.0 1.15 2.47 82.6 2.28 Brown and
labrosus) percent protein) seaweed meal Camilleri
(1997)
Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review
TABLE 1.2 (cont.)
Algae

Performance of various fish species fed fresh algae or dried algal meal
Algae/ fish species Rearing system Rearing Control diet Composition of test diet Inclusion Fish SGR SGR as FCR References
days level size (percent) percent
(percent) (g) of
control
Porphyra sp./ Red seabream Flow-through 42 Fishmeal based 5 percent Porphyra 5.0 15.4 3.47 111.6 1.52 Kalla et al.
(Pagrus major) system semi-purified spheroplasts added to diet (2008)
diet (51 percent
protein)
Ulva rigida/ European seabass Recirculation 70 Fish protein 5 and 10 percent fish 5.0 4.7 2.63 89.8 1.68 Valente et
(Dicentrarchus labrax) system hydrolysate based protein hydrolysate replaced 10.0 4.7 2.54 86.7 1.80 al. (2006)
diet (60.8 percent by dried seaweed
protein)
Gracilaria cornea/ European Recirculation 70 Fish protein 5 and 10 percent fish 5.0 4.7 2.63 89.8 1.74 Valente et
seabass (Dicentrarchus labrax) system hydrolysate based protein hydrolysate replaced 10.0 4.7 1.78 60.8 2.31 al. (2006)
diet (60.8 percent by dried seaweed
protein)
Gracilaria busra-pastonis/ Recirculating 70 Fish protein 5 and 10 percent fish 5.0 4.7 2.98 101.7 1.56 Valente et
European seabass system hydrolysate based protein hydrolysate replaced 10.0 4.7 3.37 115.0 1.48 al. (2006)
(Dicentrarchus labrax) diet (60.8 percent by dried seaweed
protein)
Gracilaria lichenoides/ rabbitfish Floating net 100 Carp starter Fresh live seaweed was fed 100.0 50.1 Negative growth displayed. Tacon et al.
(Siganus canaliculatus) cages pellet (27 percent as sole diet SGR of control 0.63 percent (1990)
protein)
Eucheuma cottonii/ rabbitfish Floating net 100 Carp starter Fresh live seaweed was fed 100.0 48.8 Negative growth displayed. Tacon et al.
(Siganus canaliculatus) cages pellet (27 percent as sole diet SGR of control 0.63 percent (1990)
protein)
Gracilaria sp./ Giant tiger Brackishwater 60 Soybean and 1, 2, 3 and 6 percent protein 5.0 0.024 7.88 98.3 3.33 Briggs and
prawns (Penaeus monodon) recirculatory fish meal based of control feed replaced by 10.0 8.03 100.1 3.35 Funge-
system diet (35 percent seaweed meal. Seaweed Smith
protein) meal incorporated by 15.0 7.88 98.3 3.50 (1996)
replacing soybean meal and 30.0 7.33 91.4 4.14
wheat flour
11
12

Table 1.3
Results of investigations on the use of algae as additives in fish feed
Algae1 Inclusion Fish species Effect References
level (percent)
Blue-green algae
Spirulina 2.0 Red sea bream Improved carcass quality through modification of muscle lipids Mustafa et al. (1994a)
2.0 Red sea bream Improved muscle quality; increased firmness and robustness of raw meat; and improved Mustafa, Umino and
growth and protein synthetic activity Nakagawa (1994)
5.0 Red sea bream Elevated growth rates; improved feed conversion, protein efficiency and muscle protein Mustafa et al. (1994b)
deposition
5.0 Nibbler Improved growth Nakazoe et al. (1986)
… Striped jack Improved flesh texture and taste Liao et al. (1990)
2.5 Cherry salmon Elevated growth rates, bright skin colour and fin appearance; improved flavour and firm Hensen (1990)
flesh
0.5 Yellowtail Increased survivability and improved weight gain Hensen (1990)
Spirulina maxima 20.9 [40.0 Mozambique Final body weight, daily weight gain, SGR, feed intake, PER and apparent nitrogen Olvera-Novoa et al.
replacement tilapia utilization showed no significant differences with control diet (1998)
of fish meal]
Brown algae
Ascophyllum 5.0 & 10.0 Red sea bream Improved growth and feed efficiency at 5 percent inclusion level Yone, Furuichi and
nodosum Urano (1986a)
5.0 Red sea bream Delayed absorption of dietary carbohydrate and protein. The dietary nutrients are utilized Yone, Furuichi and
effectively by this delaying effect of the seaweed; thus the growth and feed efficiency of
Urano (1986b)
red sea bream are improved
5.0 Red sea bream Elevated growth rates; improved feed conversion, protein efficiency and muscle protein Mustafa et al. (1994b)
deposition
5.0 Red sea bream Increased growth, feed efficiency and protein deposition. Elevated liver glycogen and Mustafa et al. (1995)
triglyceride accumulation in muscle
0.5 Yellowtail Prevented a nutritional disease that causes retardation of growth and high mortality Nakagawa et al.
(1986)
Undaria pinnatifida 5.0 Rockfish Showed prominent physiological effects on haematocrit value and red blood cell number Yi and Chang (1994)
5.0 & 10.0 Red sea bream Improved growth and feed efficiency, and higher muscle lipid deposition at 5 percent level Yone, Furuichi and
of inclusion Urano (1986a)
Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review
TABLE 1.3 (cont.)
Results of investigations on the use of algae as additives in fish feed
Algae

Algae1 Inclusion Fish species Effect References


level (percent)
Red algae
Porphyra yezoensis 5.0 Red sea bream Increased growth, feed efficiency and protein deposition. Elevated liver glycogen and Mustafa et al. (1995)
triglyceride accumulation in muscle
Porphyra yezoensis 2.0 Yellowtail Improved flesh quality Morioka et al. (2008)
Porphyra spheroplasts 5.0 Red sea bream Survival, growth and nutrient retention significantly higher than control Kalla et al. (2008)
Green algae
Ulva conglobata 5.0 Nibbler Improved growth Nakazoe et al. (1986)
Ulva pertusa 2.5, 5.0, 10.0 Black sea Ulva meal diets repressed lipid accumulation in intraperitoneal body fat without loss Nakagawa et al.
& 15.0 bream of growth and feed efficiency. Fish fed 2.5, 5 and 10 percent Ulva meal did not show (1993)
significant body weight loss during wintering. During starvation, lipid reserves were
preferentially mobilized for energy
Ulva pertusa extract 10.0 Black sea Improved tolerance to hypoxia Nakagawa et al.
bream (1984)
Ulva pertusa 5.0 Red sea bream Activated lipid mobilization and suppressed protein breakdown observed during Nakagawa and
starvation for fish fed Ulva meal supplemented diet before starvation. Preferential use of Kasahara (1986)
glycogen observed
5.0 Red sea bream Demonstrated a decrease in susceptibility to Pasteurella piscicida, an elevation of Satoh, Nakagawa and
phagocytosis and spontaneous haemolytic and bactericidal activity Kasahara (1987)
5.0 Red sea bream Increased growth, feed efficiency and protein deposition. Elevated liver glycogen and Mustafa et al. (1995)
triglyceride accumulation in muscle
1
Algae were added as dried meal in all diets except otherwise stated
13
14 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

However, the conclusions of the latter authors are confused by the fact that the test
diets were not iso-nitrogenous with the control diet; in fact test diets had a lower
protein level.
Total replacement of fishmeal by algal meal showed very poor growth responses
for O. niloticus (Appler and Jauncey, 1983; Appler, 1985) and T. zillii (Appler, 1985).
Appler and Jauncey (1983) recorded a SGR of 58 percent of control diet when the
filamentous green alga (Cladophora glomerata) meal was used as the sole source of
protein for Nile tilapia. Similarly, Appler (1985) recorded SGRs of 44 percent and 56
percent of control diets when the filamentous green alga (Hydrodictyon reticulatum)
meal was used as the sole source of protein for O. niloticus and T. zillii.
Tacon et al. (1990) used fresh live seaweeds (Gracilaria lichenoides and Eucheuma
cottonii) as the total diet for rabbitfish in net cages. In both cases negative growth was
displayed, although the daily feed intake was more than the control diet. On a dry
matter basis, the daily feed intake was 1.99 and 1.98 g/fish/day respectively for
E. cottonii and G. lichenoides, while the feed intake for carp pellets (control diet)
was 1.80 g/fish/day. Apparently, a good feeding response was observed for both the
seaweeds but very poor feed efficiency was displayed. Apart from commonly observed
impaired growth, the use of algae as the sole source of protein in fish feed can also
result in malformation (Meske and Pfeffer, 1978).
The apparently poor performance of fish fed diets containing higher inclusion
levels of algae may be attributable to several factors. Appler (1985) observed that most
of the aquatic plants including algae contain 40 percent or more of carbohydrate, of
which only a small fraction consists of mono- and di-saccharides. Low digestibility
of plant materials has been attributed to a preponderance of complex and structural
carbohydrates. The poor digestibility and the subsequent poor levels of utilization
obtained for both tilapia species with increased dietary algal levels may thus be
attributable in part to the presence of indigestible algal materials. Pantastico, Baldia and
Reyes (1985) reported that newly hatched Nile tilapia fry (mean weight 0.7 mg) did not
survive at all when unialgal cultures of Euglena elongata and Chlorella ellipsoidea were
fed to them. These authors concluded that the mortality of tilapia fry might be due to
factors such as toxicity and cell-wall composition of the algae fed. This phenomenon
might also be attributed to poor digestion of plant material by the less developed
digestive system of newly hatched larva. In contrast, Chow and Woo (1990) recorded
significantly higher gut cellulase activity in O. mossambicus fed Spirulina, indicating the
ability of this tilapia species to digest cellulose, the main constituent of plant cell walls.
Ayyappan et al. (1991) conducted a Spirulina feeding experiment with carp species. The
fry stage of catla (Catla catla), rohu (Labeo rohita), mrigal (Cirrhinus mrigala), silver
carp (Hypophthalmicthys molitrix), grass carp (Ctenopharyngodon idella) and common
carp (Cyprinus carpio) were fed with an experimental diet in which 10 percent dried
Spirulina powder was added to a 45:45 mixture of rice bran and groundnut oil cake. A
50:50 bran-groundnut oil cake control diet was used. The mean specific growth rates
of fish fed on the two diets were: catla 0.17, 0.27; rohu 0.19, 0.63; mrigal 0.54, 0.73;
grass carp 0.02, 0.40; and common carp 0.15, 0.20; with significant differences between
the treatments (F1,4 = 8.88; P < 0.05) and fish species (F4,4 = 5.03; P < 0.10). Rohu and
mrigal showed significantly (P < 0.05) higher SGRs than catla and common carp. These
results clearly demonstrated the beneficial effect of the Spirulina diet on the yield and
quality of carp fry.
Dietary supplementation of Chlorella ellipsoidea powder at 2 percent on a dry-
weight basis showed higher weight gain and improved feed efficiency and protein
efficiency ratios in juvenile Japanese flounders (Paralichthys olivaceus); the addition of
Chlorella had positive effects as it significantly reduced serum cholesterol and body fat
levels and also led to improved lipid metabolism (Kim et al., 2002).
Algae 15

Clearly, no definite conclusions can be arrived at this stage about the value of using
macroalgae as major dietary ingredients or protein sources in aquafeeds. Moderate
growth responses and good food utilization (FCR 1.5–2.0) were generally recorded
when dried algal meal were used as a partial replacement of fishmeal protein. However,
the collection, drying and pelletization of algae require considerable time and effort.
Furthermore, cultivation costs would have to be taken into consideration. Therefore,
further cost-benefit on-farm trials that take these costs into consideration are needed
before any definite conclusions on the future application of algae as fish feed can be
drawn.

1.5.2 Algae as feed additives


The main applications of mi­croalgae for aquaculture are associated with nutrition,
being used fresh (as sole component or as food additive to basic nutrients) for
colouring the flesh of salmonids and for induc­ing other biological activities (Muller-
Feuga, 2004). Several investigations have been carried out on the use of algae as additives
in fish feed. Feeding trials were carried out with many fish species, most commonly
red sea bream (Pagrus major), ayu (Plecoglossus altivelis), nibbler (Girella punctata),
striped jack (Pseudoceranx dentex), cherry salmon (Oncorhynchus masou), yellowtail
(Seriola quinqueradiata), black sea bream (Acanthopagrus schlegeli), rainbow trout
(Oncorhynchus mykiss), rockfish (Sebastes schlegeli) and Japanese flounder (Paralichthys
olivaceus). Various types of algae were used; the most extensively studied ones have been
the blue-green algae Spirulina and Chlorella; the brown algae Ascophyllum, Laminaria
and Undaria; the red alga Porphyra; and the green alga Ulva. Fagbenro (1990) predicted
that the incidence of cellulase activity could be responsible for the capacity of the
catfish Clarias isherencies to digest large quantities of Cyanophyceae.
A summary of the results of selected feeding trials with algae as feed additives is
presented in Table 1.3. Most of these research studies were conducted in Japan with
Japanese fish species, although the results may well be applicable to other species and
in other countries.
Table 1.3 shows that dried algal meals or their extracts have been added to test fish diets
at levels up to 21 percent level. The responses of test fish fed algae supplemented diets
were compared with fish fed standard control diets. Although various types of algae and
fish species were used in these evaluations, not all algae were evaluated as feed additives
for every different species. As the main biochemical constituents and digestibility are
different among algae, the effect of dietary algae varies with the algae and fish species
(Mustafa and Nakagawa, 1995). While studying the effect of two seaweeds (Undaria
pinnatifida and Ascophyllum nodosum) at different supplementation levels for red sea
bream, Yone, Furuichi and Urano (1986a) observed best growth and feed efficiency from
a diet containing 5 percent U. pinnatifida followed by a diet containing 5 percent A.
nodosum. Similarly, Mustafa et al. (1994b) observed more pronounced effects on growth
and feed utilization of red sea bream by feeding a diet containing Spirulina compared
to one containing Ascophyllum. In another study, Mustafa et al. (1995) studied the
comparative efficacy of three different algae (Ascophyllum nodosum, Porphyra yezoensis
and Ulva pertusa) for red sea bream and noted that feeding Porphyra showed the most
pronounced effects on growth and energy accumulation, followed by Ascophyllum and
Ulva. However, research results obtained so far do not specifically identify any specific
algae as the most suitable as feed additives for any particular fish species.
Nevertheless, the results of various research studies show that algae as dietary
additives contribute to an increase in growth and feed utilization of cultured fish due
to efficacious assimilation of dietary protein, improvement in physiological activity,
stress response, starvation tolerance, disease resistance and carcass quality. In fish fed
algae-supplemented diets, accumulation of lipid reserves was generally well controlled
and the reserved lipids were mobilized to energy prior to muscle protein degradation
16 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

in response to energy requirements. In complete pelleted diets, algal supplementation


of 5 percent or less was found to be adequate.
Spirulina are widely used as feed additives in the Japanese fish farming industry.
Henson (1990) reported that Spirulina improved the performances of ayu, cherry
salmon, sea bream, mackerel, yellowtail and koi carp. The levels of supplementation
used by Japanese farmers are 0.5-2.5 percent. Henson (1990) further reported that
Japanese fish farmers used about US$2.5 million worth of Spirulina in 1989. Five
important benefits reported by using a feed containing this alga were improved
growth rates; improved carcass quality and colouration; higher survival rates; reduced
requirement for medication; and reduced wastes in effluents. However, the high cost
of most of these algae may limit their use to the commercial production of high value
fish only.
17

2. Floating aquatic macrophytes


– Azolla

Floating aquatic macrophytes are defined as plants that float on the water surface,
usually with submerged roots. Floating species are generally not dependent on soil or
water depth.
Azolla spp. are heterosporous free-floating freshwater ferns that live symbiotically
with Anabaena azollae, a nitrogen-fixing blue-green algae. These plants have been
of particular interest to botanists and Asian agronomists because of their association
with blue-green algae and their rapid growth in nitrogen deficient habitats (Islam and
Haque, 1986). The genus Azolla includes six species distributed widely throughout
temperate, sub-tropical and tropical regions of the world. It is not clear whether the
symbiont is the same in the various Azolla species.
Azolla spp. consists of a main stem growing at the surface of the water, with
alternate leaves and adventitious roots at regular intervals along the stem. Secondary
stems develop at the axil of certain leaves. Azolla fronds are triangular or polygonal and
float on the water surface individually or in mats. At first glance, their gross appearance
is little like what are conventionally thought of as ferns; indeed, one common name for
them is duckweed ferns. Plant diameter ranges from 1/3 to 1 inch (1-2.5 cm) for small
species like Azolla pinnata to 6 inches (15 cm) or more for A. nilotica (Ferentinos,
Smith and Valenzuela, 2002).

2.1 Classification
The genus Azolla belongs to the single genus family Azollaceae. The six recognizable
species within the genus are grouped under two subgenera: Euazolla and
Rhizosperma.
The four species under the sub-genus Euazolla are A. filiculoides, A. caroliniana,
A. mexicana and A. microphylla. It is thought that these four species originated from
temperate, sub-tropical and tropical regions of North and South America (Van Hove,
1989). However, Zimmerman et al. (1991) found three of these species (A. caroliniana,
A. mexicana and A. microphylla) to have close taxonomic affinity and similar responses
to phosphorus deficiency and recommended that they be considered as a single
species.
The two species under sub-genus Rhizosperma are A. nilotica and A. pinnata. A.
nilotica is a native of East Africa and can be found from Sudan to Mozambique (Van
Hove, 1989). A. pinnata has two different varieties that vary in their distribution
patterns. A. pinnata var. imbricata originates from subtropical and tropical Asia while
A. pinnata var. pinnata occurs in Africa and is known as African strain.

2.2 Characteristics
2.2.1 Importance
Because Azolla has a higher crude protein content (ranging from 19 to 30 percent)
than most green forage crops and aquatic macrophytes and a rather favourable
essential amino acid (EAA) composition for animal nutrition (rich in lysine), it has
also attracted the attention of livestock, poultry and fish farmers (Cagauan and Pullin,
1991). In Asia Azolla has been long used as green manure for crop production and
a supplement to diets for pigs and poultry. Some strains of Azolla can fix as much
as 2-3 kg of nitrogen/ha/day. Azolla doubles its biomass in 3-10 days, depending
18 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

on conditions, and easily reaches a standing crop of 8-10 tonnes/ha fresh weight in
Asian rice fields; 37.8 tonnes/ha fresh weight (2.78 tonnes/ha dry weight) has been
reported for A. pinnata in India (Pullin and Almazan, 1983). Recently, Liu et al. (2008)
have reported the application of Azolla as a controlled ecological life support system
(CELSS) for its strong photosynthetic oxygen-releasing capacity. Azolla provides a
protected environment and a fixed source of carbon to the blue-green filamentous algae
Anabaena spp. (Wagner, 1997).

2.2.2 Environmental requirements


The potential for rearing Azolla is restricted by climatic factors, water and inoculum
availability, the incidence of pests, phosphorus requirements and the need for
labour intensive management (Cagauan and Pullin, 1991). Water is the fundamental
requirement for the growth and multiplication of Azolla. The plant is extremely
sensitive to lack of water. Although Azolla can grow on wet mud surfaces or wet pit
litters, it prefers growing in a free-floating state (Becking, 1979). A strip of water not
more than a few centimetres deep favours growth because it provides good mineral
nutrition, with the roots not too far from the soil, and also because it reduces wind
effects (Van Hove, 1989). Strong winds can accumulate Azolla to one side of the stretch
of water, creating an overcrowded condition and thus slowing growth.
Azolla can survive a water pH ranging from 3.5-10, reported optimum growth
occurring at pH 4.5-7.0. Watanabe et al. (1977) reported that the growth of Azolla was
optimum at pH 5.5 and FAO (1977) recorded that soils of pH 6 to 7 support the best
growth.
Salinity tolerance of Azolla species also varies. The growth rate of A. pinnata
declines as its salinity increases above 380 mg/l (Thuyet and Tuan, 1973). According
to Reddy et al. (2005) Azolla can withstand salinity of up to 10 ppt but Haller, Sutton
and Burlowe (1974) reported that the growth of A. caroliniana ceases at about 1.3 ppt
and higher concentrations result in death. A. filiculoides has been reported to be most
salt-tolerant (I. Watanabe pers. comm., cited by Cagauan and Pullin, 1991).
Azolla grows in full to partial shade (100-50 percent sunlight) with growth
decreasing quickly under heavy shade (Ferentinos et al., 2002). Generally, Azolla
requires 25-50 percent full sunlight for its normal growth; slight shade is of benefit
to its growth in field conditions. The biomass production greatly decreases at a light
intensity lower than 1 500 lux (Liu et al., 2008).
Like all other plants, Azolla needs all the macro- and micro-nutrients for its normal
growth and vegetative multiplication. All elements are essential; phosphorus is often
the most limiting element for its growth. For normal growth, 0.06 mg/l/day is required.
This level can be achieved in field conditions by the split application of superphosphate
at 10-15 kg/ha (Sherief and James, 1994). 20 mg/l is the optimum concentration
(Ferentinos et al., 2002). The symptoms of phosphorous deficiency are red-coloured
fronds (due the presence of the pigment anthocyanin), decreased growth and curled
roots. Macronutrients such as P, K, Ca and Mg and micronutrients such as Fe, Mo and
Co have been shown to be essential for the growth and nitrogen fixation of Azolla
(Khan and Haque, 1991).
The temperature tolerance of Azolla varies widely between its species and strains. In
general, Azolla has low tolerance to high temperature and that restricts its use in tropical
agriculture. There are, however, strains that can adapt successfully to high temperature.
Cagauan and Pullin (1991) ranked different Azolla species from the most to the least
heat-tolerant, based on the optimum temperature for good growth: A. mexicana > A.
pinnata var. pinnata > A. microphylla > A. pinnata var. imbricata, A. caroliniana > A.
filiculoides (Table 2.1). In general, the optimum temperature for growth of all Azolla
species is around 25 ºC, except that of A. mexicana, whose optimum temperature is
Floating aquatic macrophytes – Azolla 19

Table 2.1
Temperature tolerance of five species of Azolla
Subgenera Species Water temperature (ºC)
Minimum Maximum Optimum for growth
Euazolla A. filiculoides 0-10 38-42 20-25
A. caroliniana <0-10 45 20-30
A. mexicana - - 30-33
A. microphylla 5-8 45 25-30
Rhizosperma A. pinnata
A. pinnata var. <5 >40 16-33
pinnata
A. pinnata var. 0 45 20-30
imbricata
Source: modified from Cagauan and Pullin (1991)

~30 ºC. According to Sherief and James (1994), the favourable water temperature for
rapid multiplication of Azolla is generally between 18 and 26 ºC.
The optimum relative humidity for Azolla growth is between 85-90 percent. Azolla
becomes dry and fragile at a relative humidity lower than 60 percent.

2.3 Production
Multiplication of Azolla in nature and in the laboratory is entirely through vegetative
reproduction. However, sexual reproduction, which is essential to the survival of the
population during temporary adverse conditions also, occurs. When Azolla fronds
reach a certain size depending on the species and the environment, generally 1 to 2 cm
in diameter, the older secondary stems detach themselves from the main stem as a result
of the formation of an abscission layer, thus giving rise to new fronds. This is the most
usual mode of multiplication.
Sherief and James (1994) have described a simple Azolla nursery method for its
large-scale multiplication in the field for Indian farmers. The field for an Azolla nursery
must be thoroughly prepared and levelled uniformly. It is divided into different plots
by providing suitable bunds and irrigation channels. Water is manipulated at a depth
of 10 cm. Ten kg of fresh cattle dung mixed in 20 L of water is sprinkled in each plot
and an Azolla inoculum of 8 kg is introduced to each plot. Superphosphate (100 g)
is applied in three split doses at intervals of four days as a top dressing fertilizer. For
insect control, furadone granules at 100 g/plot are applied seven days after inoculation.
Fifteen days after inoculation, Azolla is harvested. From one harvest, 40-55 kg of fresh
Azolla is obtained from each plot. Reddy and DeBusk (1985) reported the yield of
Azolla (A. caroliniana) to be 10.6 t DM/ha/year in nutrient non-limiting waters of
central Florida, USA.
According to Ferentinos et al. (2002) the nitrogen fixation capacity of Azolla was
found to vary from 53-1 000 kg/ha with a dry matter production of 39-390 tonnes/ha,
in crop cycles of 40-365 days. The linear growth phase is usually between 6 and
21 days and is characterized by low lignin and cell wall fractions. Due to its high lignin
content (20 percent), nitrogen is released slowly from the plant initially, with about
two-thirds released on the first 6 weeks after application. Under flooded conditions,
40-60 percent of the available N is released after 20 days and 55-90 percent within
40 days after application
Reddy et al. (2005) described the production of Azolla in earthen raceways
(10.0 m x 1.5 m x 0.3 m) in CIFA, Bhubaneswar. 6 kg of Azolla was inoculated in each
raceway. 50 kg single super phosphate and pesticide (1-2 mg/l) were applied and a
water depth of 5-10 cm was maintained. 18-24 kg/raceway/week was produced. About
one tonne of Azolla could be harvested every week from water spread area of 650 m2,
with a phosphorus input-nitrogen output ratio of 1:4.8.
20

Table 2.2
Chemical analyses of various Azolla species
Azolla species Moisture Composition1 Minerals Reference
(percent) (percent DM1) (percent DM)
CP EE CF Ash CC Ca P K
A. filiculoides 93.52 25.0-28.5 3.1 n.s. 17.3 4.4-11.5 0.5-1.5 1.0-1.5 6.0 modified from Cagauan and
Pullin (1991)
A. caroliniana 20.6-22.6 n.s. n.s. n.s. 8.5 0.6 1.3 5.3 modified from Cagauan and
Pullin (1991)
A. pinnata var. imbricate 26.0 n.s. n.s. n.s. 4.1 0.4 1.3 4.5 modified from Cagauan and
Pullin (1991)
A. pinnata (tank culture) 18.2 1.3 n.s. 21.7 n.s. 1.6 0.6 n.s. modified from Cagauan and
Pullin (1991)
A. pinnata (field culture) 22.2 2.9 n.s. 18.3 14.7 n.s. n.s. n.s. modified from Cagauan and
Pullin (1991)
A. pinnata 21.4 2.7 12.7 16.2 Alalade and Iyayi (2006)
A. microphylla (lab. culture) 21.8 2.9 n.s. 21.6 15.6 n.s. n.s. n.s. modified from Cagauan and
Pullin (1991)
A. microphylla (field culture) 20.0-26.0 3.0-3.5 n.s. 14-15 4.0-14.0 0.7 1.6 5.5 modified from Cagauan and
Pullin (1991)
A. microphylla hybrid (field culture) 19.0 4.0-4.5 n.s. 16.0-17.0 2.5-3.0 n.s. n.s. n.s. modified from Cagauan and
Pullin (1991)
Various Azolla spp. 13.0-30.0 4.4-6.3 n.s. 9.7-23.8 5.6-15.2 0.2-0.7 0.1-1.6 0.3-6.0 Reddy et al. (2005)
Azolla sp. n.s. n.s. n.s. n.s. n.s. 1.0 0.4 2.5 Ferentinos, Smith and Valenzuela,
(2002)
1
CP = crude protein; EE = ether extract; CC = crude cellulose; Ca = calcium; P = phosphorus; K = potassium
2
Data obtained from Tacon (1987)
Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review
Floating aquatic macrophytes – Azolla 21

2.4 Chemical composition


The chemical composition of Azolla species varies with ecotypes and with the
ecological conditions and the phase of growth. The dry matter percentage of different
Azolla species varies widely and there is little agreement between the published data on
this subject: values of 5 to 7 percent can, however, be taken as fair estimates (Van Hove,
1989). A summary of the chemical composition of various Azolla species is presented in
Table 2.2. Generally, the crude protein content is about 19-30 percent DM basis during
the optimum conditions for growth (Peters et al., 1979; Becking, 1979). Under natural
conditions, values near 20-22 percent are frequent. The protein contents of Azolla
species are comparable to or higher than that of most other aquatic macrophytes. Like
most of the other aquatic macrophytes, Azolla have high ash contents, varying between
14-20 percent. No clear interspecific difference in the crude lipid levels of various
Azolla species occurs; the value is around 3-6 percent on a DM basis.
Amino acid compositions of Azolla spp. are presented in Annex 1 Table 2. Generally,
these species are low in methionine but high in lysine (except for A. pinnata). A.
microphylla is richest in all EAA except in methionine. The poorest species with respect
to most of the EAA is A. filiculoides although lysine and methionine contents in this
species are moderate. The EAA composition of Azolla species is comparable to that of
the aquatic plants commonly used as fish feed ingredients. The lysine and methionine
contents of most Azolla species appear to be higher than some ‘conventional’ plant
protein sources.

2.5 Use as aquafeed


In spite of its attractive nutritional qualities and relative ease of production in ponds
and rice-fields, reports on the use of Azolla in aquaculture are extremely limited. The
value of Azolla as a fish feed is still being studied. Available literature on the use of
Azolla for this purpose has been reviewed as follows under the headings experimental
studies and field studies.

2.5.1 Experimental studies


A few studies have been carried out in aquaria to examine the preference for various
Azolla species by different cichlid species and a carp hybrid. These tests were carried
out using fresh Azolla; the results are summarized in Table 2.3. These preference
tests indicate that A. caroliniana (Figure 2.1) is one of the most preferred species for
cichlids.
A number of growth studies have been carried
out to evaluate Azolla as fish feed under laboratory Figure 2.1
rearing conditions. Most of these studies were Azolla/mosquito fern (Azolla caroliniana)
conducted on cichlids and little or no attempt
was made to use Azolla as a feed for grass carp,
a predominantly macrophytophagous feeder. In
these studies, Azolla was fed either in fresh or
dried powdered form as a whole feed or by
partially replacing fishmeal in pelleted diets.
Almazan et al. (1986) conducted a study where
A. pinnata was fed to Nile tilapia (Oreochromis
niloticus) fingerlings and adult males. Fingerlings
were fed Azolla in fresh, powder, and pellet form,
replacing the complete control diet mix from 10
percent to 90 percent. The control diet consisted
of 40 percent fishmeal, 40 percent rice bran, 10
percent cornstarch, 9 percent corn meal and 1 Source: www.msrosenthal.com/Ferns/images/Florida_Images/
Azolla_caroliniana.jpg
percent Afsillin (micronutrient premix). Negative
22 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

Table 2.3
Preference of Azolla spp. by different fish species
Fish species Preferred Azolla species1 Reference
Cichlasoma fenestratum (1) A. microphylla Antoine et al. (1986)
(2) A. caroliniana
Oreochromis niloticus (1) A. filiculoides Antoine et al. (1986)
(2) A. microphylla
(3) A. caroliniana
O. mossambicus A. caroliniana Lahser (1967)
O. niloticus A. microphylla Fiogbé, Micha and Van Hove
(2004)
Tilapia rendalli (1) A. caroliniana Micha et al. (1988)
(2) A. pinnata var. pinnata
(3) A. microphylla
(4) A. filiculoides
Hybrid carp (grass carp x bighead carp) A. caroliniana Cassani (1981)
1
Azolla species are arranged chronologically for each fish species in order of preference i.e. from most preferred to less preferred

or very slow growth was obtained in all Azolla-incorporated diets. A lowering of growth
performance and FCRs was observed with increasing Azolla incorporation. Adult male
tilapia were fed dried Azolla pellets or fresh Azolla ad libitum. Despite feeding to
satiation, tilapia suffered weight loss in a 30-day feeding trial. The experiments were
carried out in aquaria. Similarly, Antoine et al. (1986) working with O. niloticus and
Cichlasoma melanurum and Micha et al. (1988) with O. niloticus and Tilapia rendalli
reported poor growth and feed utilization when they were fed A. microphylla-based
diets. Antoine et al. (1986) and Micha et al. (1988) conducted a 70-day growth trial and
fed the target species with three different diets: commercial pellets (34 percent protein);
fresh Azolla plus 28 percent protein commercial pellets (50:50); and fresh Azolla
(22 percent protein).
In other studies, El-Sayed (1992; 2008) reported extremely poor performance for
O. niloticus fingerlings and adults fed diet based on A. pinnata. This author incorporated
dried Azolla powder at 25, 50, 75 and 100 percent replacement of fishmeal protein in
a fishmeal-based control diet in a 70-day trial. Fresh Azolla as a total diet was also
used as a control. Growth and feed utilization efficiency of fish fed with the control
diet was significantly higher compared to fish fed with Azolla-supplemented diets.
The performance of fish was inversely related to the increasing dietary incorporation
of Azolla. In fish fed with the total Azolla (dry/fresh) based diet, the reduction was
extremely sharp. Fresh Azolla-fed adults started losing weight from the 7th week. Fish
fed with fresh plant material had significantly higher moisture content than fish fed
with formulated diets. Body protein and lipid levels were negatively correlated with the
concentrations of Azolla in the diets; ash content showed a positive correlation.
In all the experimental studies examined above (Almazan et al., 1986; Antoine et
al., 1986; Micha et al., 1988), fish growth was higher in the conventional control diets
than in the Azolla-based diets. Fish died or negative growth was recorded when fed
exclusively with fresh Azolla.
In apparent contrast, Santiago et al. (1988) found that O. niloticus fry fed rations
containing up to 42 percent of A. pinnata outperformed fish fed a fishmeal-based
control diet. Growth and feed utilization of O. niloticus fry improved with increased
dietary inclusion of Azolla and the survival was unaffected. These results differed from
the studies of Almazan et al. (1986), Antoine et al. (1986) and Micha et al. (1988) and it
must be pointed out that Santiago et al. (1988) used a 35 percent protein diet with early
fry (11-14 mg). In the other studies, the crude protein level was generally lower and the
studies were carried out with advanced fry, fingerling or adults. El-Sayed (2008) noted
that young Nile tilapia utilized Azolla more efficiently than adults.
Fiogbé, Micha and Van Hove (2004) obtained quite favourable results with
Azolla-based diets fed to juvenile Oreochromis niloticus grown in a recirculating
system. Six diets were formulated with almost isonitrogenous levels of protein
Floating aquatic macrophytes – Azolla 23

(27.25-27.52 percent DM) but different levels of dry Azolla meal (0, 15, 20, 30, 40
and 45 percent). All diets with incorporated Azolla meal showed weight gain. The
Azolla-free diet and the diet containing 15 percent Azolla produced the same growth
performance. The least expensive diet, which contained 45 percent Azolla, also
showed growth and was thought to be potentially useful as a complementary diet for
tilapia raised in fertilized ponds. These authors noted that mixing Azolla with some
agricultural by-products such as rice bran; the use of fermentable by-products such as
yeasts; or the addition of purified enzymes; might improve ingestion and digestibility.
Carcass compositions of fish were reported to be markedly affected by feeding
with Azolla. Antoine et al. (1986) observed that when fed with fresh Azolla, both
O. niloticus and C. melanurum had higher moisture and lower lipid concentrations.
Similar results and an increase in carcass ash content for O. niloticus and T. rendalli
were reported by Micha et al. (1988). El-Sayed (1992) also made similar observations
when he fed fresh and dried A. pinnata to O. niloticus. However, his observation differs
from the previous authors to the extent that carcass protein content was negatively
correlated with Azolla levels in the diets, while the other workers recorded no effect
on carcass protein content.
The poor growth of fish fed with diets containing higher levels of Azolla may be due
to excesses or deficiencies of amino acids, according to Fiogbé, Micha and Van Hove
(2004). Cole and Van Lunen (1994) found that inadequate levels of essential amino
acids resulted in depression of food intake and growth. Deficiencies of one or more
amino acids are known to limit protein synthesis, growth or both.

2.5.2 Field studies


Until recently, reports of on-farm utilization of Azolla were limited (Cagauan and
Pullin, 1991). At that time reports came only from China and Vietnam (Figure 2.2).
More recently Azolla has increasingly been used as feed and/or fertilizer in studies with
rice-fish culture systems in many other Asian countries. Reddy et al. (2005) reported
that the manuring schedule can be reduced by 30-35 percent through Anabaena azollae
—Azolla biofertilization in aquacultural ponds.

Azolla in cage culture


Pantastico, Baldia and Reyes (1986) used Figure 2.2
Harvest of fish from a pond (Hoa Binh
fresh whole A. pinnata as a supplemental
Province, Viet Nam)
feed for the cage culture of Nile tilapia in
Laguna de Bay, Philippines. Azolla was
propagated in fine mesh net enclosures in
the lake and harvested for feeding to tilapia
in cages. Four separate experiments were
conducted and weight gain was compared
with an unfed control. It was assumed that
in control cages fish grew by feeding natural
food (i.e. plankton) available in the cage. A
summary of the results is given in Table
2.4. Although higher weight gain of fish
was observed over the unfed control, the
difference in mean weight between fish fed
fresh Azolla and unfed control was about
5-10 g. The results of this cage culture study These low-input aquaculture ponds are generally stocked with
macrophytophagous fish (primarliy carp species) and fresh Azolla (Azolla
do not justify fish culture in cages using pinnata) are commonly used as supplemental feed.

Azolla as the only feed. Courtesy of M.G. Kibria


24 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

Table 2.4
Cage culture of Nile tilapia using Azolla as feed
Initial Stocking Duration Feeding rate Fresh Azolla Unfed control
weight density (months) (percent) Harvest weight (g) Harvest weight (g)
(g) (Numbers/m3)

1.3 25 6 35 and 70 30.3 and 36.3 24.7


1.6 50 4 30 75.0 64.1
6.5 100 5 20 20.2 10.9
13.5 150 3 15 29.3 20.2

Rice-fish-Azolla integration
One of the most successful uses of Azolla is its use as fertilizer and/or feed in an
integrated rice-fish-Azolla system. This system is based on convenient layout of the
fields, which allows the simultaneous development of rice, Azolla and different fish
with complementary nutritional requirements (Van Hove, 1989). In this ecological
agricultural layout, each of the partners contributes to the equilibrium of the system.
The fish (a correct mixture of planktophages, macrophytophages and polypages) derive
a benefit from Azolla - more or less, depending on the species; their waste promotes
the proliferation of plankton that is consumed by some of the fish and fertilizes the
rice. The polyphagous fish protect the rice and Azolla from a number of insects and
molluscan pests.
Cagauan and Pullin (1991) reviewed the rice-fish-Azolla integrated system and
described its physical set-up, which is provided with pits (pond refuse/ main channel)
and ditches (trenches). Lateral or peripheral trenches are interconnected with each other
and with the main channel. Trenches serve as links for the fish from the main channel
to rice field area and also as a growing area for Azolla during the paddy cultivation
period. Depending on the size of the rice field, trenches may be dug at 15-20 m
intervals in single or rib-shaped patterns. In India, a 0.2 ha rice field was provided with
0.5 m deep and 0.5 m wide trenches and a 1.0 m deep and 1.5 m wide main channel
(Shanmugasundaram and Ravi, 1992). Cagauan (1994) used 1 m wide and 0.75 m deep
pond refuge connected to an outer trench that was 0.3-0.4 m wide and 0.2-0.3 m deep
in a 200 m2 paddy field. The trenches and main channels should occupy about
10-15 percent of the rice field area (Cagauan and Pullin, 1991; Shanmugasundaram
and Balusamy, 1993). Inoculation of the rice field with Azolla at the rate
of 4.5-6.0 tonnes/ha is done 20 days before rice transplanting. Propagated Azolla
biomass is ploughed under, together with inorganic fertilizer, before rice transplanting.
The field is then reflooded to allow the Azolla that floated during the incorporation to
grow and serve as a fish fodder. In case of insufficiency of Azolla in the channels and
trenches, additional supplemental feed is given. The fish species cultured in these rice-
fish-Azolla systems are mainly Nile tilapia. Other species are common carp, Indian
major carp, Java barb, etc. Grass carp may not be a suitable species for this system, as
they may damage the rice crop by feeding on its leaves.
The use of Azolla (A. microphylla) as a fertilizer in rice-fish farming was studied
by Cagauan and Nerona (1986) and Cagauan (1994). Cagauan and Nerona (1986)
used three fertilizer regimes: Azolla only; inorganic fertilizers (urea and ammonium
phosphate) only; and Azolla plus inorganic fertilizers for rice-fish culture with Nile
tilapia as the target species. When a combination of Azolla and inorganic fertilizers was
used, it was possible to reduce the standard rate of inorganic fertilizers by half. Fish
yields were the same with Azolla or inorganic fertilizers alone, whereas the yields of
both fish and rice were higher in the combined Azolla and inorganic fertilizer regime
(Table 2.5).
Shanmugasundaram and Ravi (1992) reported the use of Azolla (A. microphylla) as
nitrogen fixing fertilizer and feed for Nile tilapia (O. niloticus) stocked in a low-lying
ricefield (0.2 ha) in the Tanjore Deltaic Zone, Tamil Nadu, India. The ricefield was
Floating aquatic macrophytes – Azolla 25

Table 2.5
Use of A. microphylla as fertilizer in rice-fish culture system- fish species (O. niloticus)
Initial Fish density Duration Fertilizer Fertilizer Quantity Fish Rice
weight (Numbers/ (days) regimes rate of N yield yield
(g) ha) (kg/ha) (kg/ha) (kg/ha) (kg/ha)

8.9-9.4 5 000 75 Azolla 3 750 5.63 45.1 2 567


only
Inorganic 150 38.5 45.0 3 096
fertilizer
Azolla 3 750 5.6
Inorganic 75 19.3 79.0 3 524
fertilizer

Source: Cagauan and Nerona (1986)

provided with trenches and connected to a main channel and the fish were raised in
these trenches. The stocking density used was 6 000/ha for fingerlings weighing 19 g.
Both fresh and dried Azolla were fed. Dried Azolla was incorporated in a supplemental
fish feed and applied at 5 percent BW/day. The formula of this supplemental feed
was stated to be Azolla (50 percent), rice bran (15 percent), chicken manure (10
percent), corn meal (5 percent), sorghum meal (5 percent), broken rice (2.5 percent)
and groundnut cake (2.5 percent). The provision of water space for the fish lowers rice
yields by about 300 kg/ha but the fish harvest compensates. Rice and fish culture yields
a net income of US$258/crop/ha, compared to US$207/crop/ha for rice alone.
Furthermore, Shanmugasundaram and Balusamy (1993) reported the use of Azolla
(A. microphylla) as feed to raise Indian major carps (catla, rohu and mrigal) stocked in
low-lying wetlands in Bhavanisagar, Tamil Nadu, India. These authors used a 0.25 ha
ricefield provided with trenches (1.0 m depth and width) to shelter the fish. Stocking
density was 3 000/ha, using a 1:1:1 ratio of catla, rohu and mrigal. Azolla was applied
twice at 2.0 tonnes/ha. Supplemental feed containing banana pseudostem and cow
dung (1:1) was fed along with rice bran at 5 percent BW/ per day. Banana pseudostem
and cow dung were incubated overnight before mixing with rice bran. Both rice
and fish yields increased, with higher benefit cost ratios (1.88) in rice-fish-Azolla
Table 2.6
Economics of rice-fish-Azolla integration in India
Treatment Rice yield Fish yield Gross return Net return Benefit cost
(kg/ha) (kg/ha) (US$/ha) (US$/ha) ratio

Rice alone 8 765 - 822 353 1.75


Rice-fish 7 813 98.5 812 297 1.57

Rice-fish-Azolla 9 226 154.0 985 463 1.88

Source: Shanmugasundaram and Balusamy (1993)

Table 2.7
Results of rice-fish-Azolla integration highlighting increase in fish yield
Culture Fish species Average harvest weight (g) Yield (tonnes/ha)
system With Azolla Without With Azolla Without
Azolla Azolla
Monoculture Oreochromis niloticus 150-200 100-150 1.20 0.63
Polyculture Cyprinus carpio 600 350 0.35 0.15
Ctenopharyngodon idella 150 130 0.17 0.15
Oreochromis niloticus 125 100 0.54 0.40
Total 1.06 0.70
26 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

integration than rice-fish cultivation (1.57) (Table 2.6). Similarly, substantial increases
in fish yield in rice-fish culture with Azolla compared to rice-fish without Azolla have
been reported by Cagauan and Pullin (1991). Fish yields from rice-Azolla-fish culture
trials were higher than those for rice-fish culture (Table 2.7). Yields from Nile tilapia in
monoculture and from polyculture of common carp, grass carp and Nile tilapia were
1.20 and 1.06 tonnes/ha/year respectively from a rice-fish-Azolla system, compared
with 0.63 and 0.70 tonnes/ha/year respectively from rice-fish fields without Azolla.
In rice farming systems, including rice-fish culture, Azolla is best incorporated as
a fertilizer during its linear growth phase, when there is maximum productivity, low
lignin content and therefore rapid decomposition. The value of Azolla as a fish feed
is also highest during its linear growth phase. The crude protein content of Azolla
is generally higher during this phase. The amino acid content of Azolla increases
during the linear growth phase and falls sharply when the growth slows down with
a corresponding increase in its lignin content. Digestibility clearly decreases after
the linear growth phase with increasing lignin content (Van Hove et al., 1987). It is
therefore important to maintain an equilibrium between the population of fish and that
of Azolla, either by introducing, when necessary, a supplementary biomass of Azolla
collected elsewhere, or by harvesting the excess biomass in order to keep the Azolla
population in the linear growth phase.

Pig-duck-fish-Azolla and fish-Azolla integration


Very few reports are available on the use of Azolla as fish feed in pond culture; however,
there are reports of integrated studies. Majhi, Das and Mandal (2006) fed grass carp
(Ctenopharyngodon idella) fingerlings with finely chopped Azolla caroliniana placed
over a feeding basket under pond conditions. Azolla was well accepted by grass carp.
The final weight gain of Azolla-fed fish was significantly higher compared to the
control fish. The net profit for production of grass carp was US$0.12/m2.
Gavina (1994) studied pig-duck-fish-Azolla integration. Nile tilapia were stocked in
three earthen ponds with a uniform water depth of 50 cm. The ponds were fertilized
with a mixture of dry pig and duck manure at the rate of 500 kg/ha. After initial
manure application, the water level was increased to 80 cm in all three experimental
ponds. The ponds were stocked at three densities: 10 000/ha; 20 000/ha; and 30 000/ha.
All treatments were manured (pigs and ducks) with 100 kg fresh material/ha/day and
supplemented with fresh Azolla at 200 g/m2/week. The consumption of Azolla by fish
was not monitored. However, it was observed that the fresh Azolla were seeded at a
Table 2.8
Weight gain comparisons of Azolla-fed fish
Fish No. Initial Final Survival Culture Total SGR Azolla
of weight weight (percent) period weight (percent) feed
fish (g) (g) (days) increase coeff.
(g)
Grass carp1 30 54.7 118.7 100 112 1 920 0.69 49.0
Crucian carp1 30 75.0 110.8 100 112 1 074 0.35 31.2
Nile tilapia1 30 24.7 163.1 100 100 4 152 1.89 52.2
Silver carp1 39 96.8 92.2 76.7 112 -1 017 -0.04 0.0
Nile tilapia 25 1.67 3.23 56 30 21.84 2.20
(15 percent
inclusion)2
Nile tilapia 25 1.70 2.28 61.3 30 8.89 0.98
(45 percent
inclusion)2
Grass carp3 - 22.7 270.3 100 150 1.65
Tilapia zillii4 15 2.2 4.7 93 91 34.88 0.83
1
modified from FAO (1988 cited by Cagauan and Pullin, 1991)
2
Fiogbé, Micha and Van Hove (2004)
3
Majhi, Das and Mandal (2006)
4
Abdel-Tawwab (2008)
Floating aquatic macrophytes – Azolla 27

rate of 200 g/m2/week (10 kg/50 m2) and cleared by fish after 6 or 7 days. It was found
that Azolla could be a viable source of supplementary feed, considering the high cost
of commercial feeds. The study was conducted for a period of three months. Mean
net yield varied between 8.22 and 10.97 kg/ha/day (3-4 tonnes/ha/year) at stocking
densities ranging between 10-30 000/ha.
Weight gain comparisons of Azolla-fed fish were carried out by the Soil and
Fertilizer Institute of the Hunan Academy of Agricultural Sciences (FAO, 1988 cited
by Cagauan and Pullin, 1991) using grass carp, Nile tilapia, crucian carp (Carassius
auratus) and silver carp (Hypopthalmychthys molitrix) (Table 2.8). The weight gain
by Azolla-fed grass carp averaged 174 g/fish compared with 134 g/fish for Nile tilapia
and 35.8 g/fish for crucian carp. A weight decrease of 4.6 g/fish was observed for silver
carp.
29

3. Floating aquatic macrophytes


– Duckweeds

Duckweeds are small (1-15 cm) free-floating aquatic plants with worldwide distribution.
They are monocotyledons belonging to the family Lemnaceae (which is derived
from the Greek word ‘Limne’, meaning pond) and are classified as higher plants
or macrophytes, although they are often mistaken for algae and some taxonomists
consider them as being members of the Araceae. Duckweeds serve as nutrient pumps,
reduce eutrophication effects and provide oxygen from their photosynthesising
activity. Duckweeds are often seen growing in thick blanket-like mats on still nutrient-
rich fresh and slightly brackish waters. They do not survive in fast moving water
(>0.3 m/sec) or water unsheltered from the wind. They grow at water temperatures
between 6 and 33 ºC (Leng, Stambolie and Bell, 1995).

3.1 Classification
Duckweed consists of four genera: Lemna, Spirodela, Wolffia and Wolffiella. So far, 37
species belonging to the four genera have been identified from different parts of the
world. Selected species are listed in Table 3.1. Taxonomically the family is complicated
by clonal characteristics (Culley et al., 1981). The most commonly available species
belong to the three genera Lemna, Spirodela and Wolffia. Illustrations of selected
species of duckweeds are given in Figures 3.1 - 3.3. It is quite common for floating mats
of duckweeds to consist of more than one species, e.g. Lemna and Wolffia.
Lemna is the largest genera of the family Lemnaceae. Lemna is among the most
complex and confusing groups within the entire family. Landolt (1986) hypothesized
that Lemna disperna and Lemna gibba are related as progenitor-derivative species and
the former species differentiated from the latter one. Reduction of some structures such
as frond size, number of nerves and the number of ovules in Lemna disperna, along
with its narrower geographic distribution, support the hypothesis that it was derived
from Lemna gibba or from a common ancestor. Lemna disperna has a chromosome
number of 2n = 40, whereas the numbers 2n = 40, 50, 70 and 80 have been found in
Table 3.1
Classification of selected species of duckweeds
Lemna Spirodela Wolffia Wolffiella
L. gibba S. biperforata W. arrhiza W. caudate
L. disperna S. intermedia W. australiana W. denticulata
L. gibba S. oligorrhiza W. columbiana W. lingulata
L. japonica S. polyrrhiza W. microscopia W. oblonga
L. minima S. punctata W. neglecta W. rotunda
L. minor
L. minuscula
L. paucicostata
L. perpusilla
L. polyrrhiza
L. turionifera
L. trisulca
L. valdiviana
30 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

Figure 3.1
Spirodela sp.

Source: DWRP (1998)

Figure 3.2
Common duckweed, Lemna minor grown
in a pond (Phu Tho Province, Viet Nam)

Courtesy of M.G. Kibria

Figure 3.3
Lemna gibba

Source: aphotoflora.com/DevonandCornwall/page15.html
Floating aquatic macrophytes – Duckweeds 31

Lemna gibba (Crawford et al., 2005). The allozyme study supports the continued
recognization of two species and is concordant with the hypothesis that the species are
related as progenitor and derivative. The reduced morphology of Lemna disperna and
the allozyme data indicate that this species originated via dispersal of Lemna gibba or
of a common ancestor of the two species.

3.2 Characteristics
Duckweeds are adapted to a wide variety of geographic and climatic zones and are
distributed throughout the world except in regions where temperature drops below
0 ºC during part of the year. Most species are found in moderate climates of tropical
and temperate zones. In deserts and extremely wet areas, duckweeds are rare. Lemna
spp., for example are very rare in regions with high or very low precipitation and are
not found in Greenland or the Aleutian Islands (Landolt, 2006). Although many species
can survive extremes of temperature, they generally grow faster under warm and sunny
conditions (Skillicorn, Spira and Journey, 1993). Most species show prolific growth
in the tropics. Various microclimatic factors such as light intensity, salinity, regional
temperature differences can influence the distribution of Lemnaceae species (Landolt,
1986). Birds and floods often disperse duckweeds to different geographic areas.

3.2.1 Reproduction
A duckweed plant consists of a single leaf or frond with one or more roots. Most
species of duckweed multiply principally through vegetative propagation by the
formation of daughter fronds from two pockets on each side of the narrow end of the
frond (Gaigher and Short, 1986). Newly formed fronds remain attached to the mother
frond during the initial growth phase and the plants therefore appear to consist of
several fronds. Species of the genus Spirodela have the largest fronds, measuring as
much as 20 mm across, while those Wolffia species are 2 mm or less in diameter. Lemna
species are intermediate in size, being about 6-8 mm. An individual frond may produce
as many as 20 daughter fronds during its lifetime, which lasts for a period of 10 days
to several weeks. The daughter frond repeats the history of its mother frond. Some of
the duckweed species, however, reproduce by producing unisexual and monoecious
flowers and seeds. For example, L. paucicostata routinely flowers and seeds. However,
the flowers are very small and rare in many species; male and female flowers are borne
on the same plant. Each inflorescence generally consists of two male flowers and
one female, but in Wolffia, there is one male and one female. The flowers are naked
or surrounded by spathe. The fruit is a utricle and the seeds are smooth or ribbed.
Vegetative reproduction is very rapid and is usually by the formation of buds of new
fronds from the reproductive pouches (Guha, 1997).
Many species of duckweed survive at low temperatures by forming a special
starchy ‘survival’ frond known as a turion. In cold weather, the turion is formed and
sunk to the bottom of the pond where it remains dormant until warm water triggers
resumption of normal growth. Several species survive at low temperatures without
forming turions. During the winter season, the fronds are greatly reduced but remain
at the surface. Occasionally, turion-like fronds will form, but the plants continue to
slowly reproduce vegetatively. These plants are probably the best plants to utilize in
a culture system, as restocking is virtually assured. L. gibba, L. valdiviana, L. minor,
L. trisulca and L. minuscula are five duckweed species that frequently show some
growth at cold temperatures.

3.2.2 Environmental requirements


A variety of environmental factors, such as water temperature, pH and nutrient
concentration, control the growth and survivability of duckweeds. The other
environmental factors that influence the growth rates of duckweed colonies are presence
32 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

of toxins in the water, crowding by overgrowth of the colony and competition from
other plants for light and nutrients. However, the growth rate of duckweed is favoured
by organic pollutants as well as inorganic nutrients (Guha, 1997). The effect of these
various factors is summarised below.

Temperature
Temperature tolerance and optima are dependent on species and possibly even on
clones. Optimum temperature for maximum growth of most groups apparently lies
between 17.5 and 30 ºC (Culley et al., 1981; Gaigher and Short, 1986). Although
some species can tolerate near freezing temperatures, growth rate declines at low
temperature. Below 17 ºC some duckweeds show a decreasing rate of growth (Culley
et al., 1981). Most species seem to die if the water temperature rises above 35 ºC. The
effect of temperature on growth is affected by light intensity, i.e. as light increases,
growth rates increase from 10 to 30 º C.
In Bangladesh, Khondker, Islam and Nahar (1993a) reported the temperature
dependent growth of S. polyrrhiza with a maximum biomass of 76.4 g/m2 recorded
in the middle of February, after which the biomass depleted gradually with the rise
in water temperature. The water temperature in the middle of February was about
19 ºC. Islam and Khondker (1991) also obtained a high growth of S. polyrrhiza at
a temperature of 16 ºC. Furthermore, Khondker, Islam and Nahar (1993b) reported
maximum growth of S. polyrrhiza at water temperatures of 22.2-22.5 ºC in a growth
study conducted in pond water. Khondker, Islam and Makhnun (1994) reported an
inverse correlation between water temperature and the biomass of L. perpusilla when
the water temperature varied between 15 and 28 ºC. These authors also noted that the
growth of this duckweed species ceased completely at 26 ºC and above.

pH
Duckweeds are generally considered to have a wide range of tolerance for pH. They
survive well from pH 5 to 9, although some authors put their range between 3 and 10.
However, pH tolerance limits of the various species differ. Stephenson et al. (1980)
noted that duckweed display optimum growth in a medium of pH 5.0-7.0. Generally,
duckweeds grow best over the pH 6.5 to 7.5 range. A doubling of biomass in 2 to 4 days
has been demonstrated at pH levels between 7 and 8 (Culley et al., 1981). Unionized
ammonia is the preferred nitrogen substrate for duckweed. An alkaline pH shifts the
ammonium-ammonia balance toward the un-ionized state and results in the liberation
of free ammonia, which is toxic to duckweed at high concentrations (100 mg NH3/L).
Islam and Paul (1977) observed that W. arrhiza grew at a pH range of 5-10, although
the optimum pH was found to be 7-8. In Bangladesh, S. polyrrhiza has been reported
to grow best at a pH between 6.5 and 7.5 (Islam and Khondker, 1991). The range of pH
for optimum growth of S. polyrrhiza reported in India was 6.8-8.5 (Kaul and Bakaya,
1976; Gopal and Chamanlal, 1991). Khondker, Islam and Makhnun (1994) reported the
pH range of 6.9 and 7.8 to be suitable for the growth of L. perpusilla. Similarly, Van
der Does and Klink (1991) observed pH of 7.36 in a lemnid habitat in the Netherlands
supporting growth of L. perpusilla. A summary of minimum, maximum and optimum
pH of various duckweed species is presented in Table 3.2.

Conductivity
Electrolyte conductivity appears to have some effect on the growth of different species
of duckweed. Zutshi and Vass (1973) found L. gibba and L. minor growing in stagnant
waters rich in electrolyte ranging from 400-500 µS/cm. Gopal and Chamanlal (1991)
reported the maximum biomass of L. perpusilla and S. polyrrhiza from roadside
pools and ditches in India within a electrolyte conductivity range of 650-1 000
µS/cm. Khondker, Islam and Nahar (1993a) recorded the complete disappearance of
Floating aquatic macrophytes – Duckweeds 33

Table 3.2
Minimum, maximum and optimum pH of various duckweed species
Duckweed species Min Max Optimum Reference
L. minor 6.1-6.7 Hicks (1932, cited by DWRP, 1997); McLay (1976)
L. perpusilla 3.2 6.9-7.8 Landolt and Kandeler (1987); Khondker, Islam and
Makhnun (1994)
S. polyrrhiza 3.7 6.5-8.5 Gopal and Chamanlal (1991); Islam and Khondker
(1991), Kaul and Bakaya (1976); Landolt and
Kandeler (1987)
S. punctata 7.0 McLay (1976)
W. arrhiza 3.5 10.0 5-7.8 Hicks (1932, cited by DWRP, 1997); Islam and Paul
(1977); Landolt and Kandeler (1987)
W. australiana 5.0 McLay (1976)
W. columbiana 6.4-7.0 Hicks (1932, cited by DWRP, 1997)
Source: DWRP (1998)

S. polyrrhiza by the end of May when a sharp fall in conductivity and alkalinity was
observed. The electrolyte conductivity of water supporting the growth of L. perpusilla
in Bangladesh reported by Islam and Khondker (1991) and Khondker, Islam and
Makhnun (1994) were 625 µS/cm and 200-890 µS/cm, respectively. High electrolyte
conductivity (1 090 µS/cm) of water supporting the growth of L. perpusilla was also
reported by Van der Does and Klink (1991) in a lemnid habitat in the Netherlands.

Nitrogen
In general, temperature and sunlight control duckweed growth more than nutrient
concentrations in the water. At high temperatures, duckweed can grow rapidly down
to trace levels of phosphorus and nitrogen. The crude protein content of duckweed
however, seems to increase to a maximum of ~40 percent DM over the range from
trace ammonia concentrations to 7-12 mg N/L (Leng, Stambolie and Bell, 1995).
Culley et al. (1981) reported that the TKN of water should not drop below 20-30
mg/l if the optimum production and a high crude protein content of duckweed are to
be maintained.
Duckweeds prefer ammonia nitrogen (NH4-N) as a source of nitrogen and
will remove ammonia preferentially, even in the presence of relatively high nitrate
concentrations. Lüönd (1980) demonstrated that higher growth rates were attained
when nitrogen was in the NH4-N rather than the NO3-N form. In organically enriched
waters, nitrogen tends to be concentrated in the NH4-N rather than the NO3-N form at
pH levels below 9 and plant growth is equally efficient in anaerobic and aerobic waters
(Said et al., 1979). In lagoons receiving organic animal wastes, the pH seldom exceeds
8, particularly with a full duckweed cover that suppresses phytoplankton growth
(Culley et al., 1978). The plants can tolerate very high ionized ammonia (NH4-N)
concentrations but the effects of unionized ammonia (NH3-N) have not been clearly
demonstrated. Urea is a suitable fertilizer and is rapidly converted to ammonia under
normal conditions. According to the results of laboratory experiments, duckweed
tolerates concentrations of elemental N as high as 375 mg/l (Rejmánková, 1979).

Phosphorus and potassium


Phosphorus is essential for rapid growth and is a major limiting nutrient after nitrogen,
although its quantitative requirement for maximum growth is generally low. Fast
growing duckweed in nutrient rich water is a highly efficient sink for both phosphorus
and potassium; little of each, however, is required for rapid growth. Saturation of
phosphate uptake by duckweed occurs at available PO4-P concentrations of 4 to
8 mg/l. Duckweed growth is not particularly sensitive to potassium or phosphorus
once an adequate threshold has been reached. Rejmánková (1979) reported good
growth of duckweed within the P concentrations of 6 to 154 mg/l. Culley et al. (1978),
34 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

working in dairy waste lagoons, achieved doubled production from 2 to 4 days at P


concentrations in excess of 35 mg/l. Reduced growth in some species occurs only after
P values dropped below 0.017 mg/l (Lüönd, 1980). Khondker, Islam and Makhnun
(1994) observed that both phosphate and silicate concentrations had significant positive
correlation with the biomass of L. perpusilla in Bangladesh.

Other minerals
A range of other important mineral levels found in water supporting Lemnaceae
is presented in Table 3.3. Although these minerals are essential for their survival,
duckweed growth is not particularly sensitive to potassium or phosphorus once an
adequate threshold has been reached.
Table 3.3
Range of important mineral contents (mg/l) of water supporting Lemnaceae
Parameter Absolute range Range of 95 percent of the samples
K 0.5 – 100 1.0 – 30

Ca 0.1 – 365 1.0 – 80
Mg 0.1– 230 0.5 – 50
Na 1.3 – >1 000 2.5 – 300
HCO3 8 – 500 10.0 – 200
Cl 0.1– 4 650 1.0 – 2 000
S 0.03 – 350 1.0 – 200
Source: modified from Landolt (1986)

In summary
Maximum, minimum and optimum requirements of some of the most important
environmental parameters (temperature, pH, conductivity, nitrogen and phosphorus)
are given in Table 3.4. It is apparent that the duckweeds are robust in terms of
survival, but sensitive in terms of thriving. They have extreme range of tolerance for
temperature, pH, conductivity, nitrogen and phosphorus with well-defined range of
optimum requirement.

3.3 Production
3.3.1 Background information
Duckweed growth is largely a function of available nutrients, temperature, light, and
degree of crowding. The highest growth rate reported for Lemnaceae under optimal
laboratory conditions is about 0.66 generations per day, which corresponds to a
doubling time of 16 hours (DWRP, 1997). Duckweeds generally double their mass in
16 hours to 2 days under optimal nutrient availability, sunlight, and water temperature.
An individual plant, a small leaflet (frond), produces 10 to 20 daughter fronds during
its lifetime, which lasts for a period of 10 days to several weeks. The daughter frond
repeats the history of its mother frond. This results in an exponential growth, at least
until the plants become crowded or run out of nutrients. Frequent periodic removal of
the plants encourages continuation of the exponential growth.
Table 3.4
Summary of some important environmental requirements of duckweed
Environmental parameters Minimum Maximum Optimum
Temperature (0C) >0 35 15-30
pH 3.0 10.0 6.5-8.0
Conductivity (µS/cm)1 200 1 090 n.s.
Nitrogen (mg/l NH3-N) Trace 375 7-12
Phosphorus (mg/l PO4-P) 0.017 154 4-8
1
Conductivity range found supporting growth of duckweed.
Floating aquatic macrophytes – Duckweeds 35

The individual clones of the same species may show quantitative variation in growth
characteristics (Rejmánková, 1975; Porath, Hepher and Koton, 1979). In the Czech
Republic, Rejmánková (1975, 1979) reported maximum dry matter yields of 3.14-
3.54 g and 7.09 g/m2/day from unmanaged fish ponds and outdoor tanks respectively,
when weekly harvesting was done. Rejmánková (1981) further reported that an
estimated annual net dry matter production of 7.5-8.0 tonnes/ha could be obtained,
provided nutrients and crowding were not limiting and harvesting was frequent.
Culley and Myers (1980) and Said et al. (1979) working in the southern USA
(9-10 months growing season) demonstrated that high nutrient lagoons and outdoor
tanks (enriched with cattle manure) yielded the dry matter equivalent of about
15 g/m2/day (55 tonnes/ha/year) when regular daily harvesting was done to remove the
excess. Said et al. (1979) reported an annual dry weight yield of 44 tonnes/ha or about
12 g/m2/day. Furthermore, Culley and Myers (1980) obtained an estimated average
annual dry matter production of 23.3 tonnes/ha with daily harvesting ranging from 10
to 35 percent of the standing crop, depending on the season. In a sewage-fed culture
system, the growth rate of Azolla spp., Spirodela spp. and Wolffia sp. were found to be
160, 350 and 280 g/m3/day, respectively (Reddy et al., 2005).
Table 3.5 presents the yields of various duckweed species under different
environmental conditions. The values varied widely, ranging from 9 to 38 tonnes
(DM)/ha/year. This wide range of productivity may be attributed to differences in
species, climatic conditions, nutrient supply and environmental conditions. Many
of the reported high yields are based on extrapolated data obtained from short-term
growth from small-scale experimental systems rather than potential long-term yields
from commercial-sized systems. Edwards (1990) reported extrapolated yields of
~20 tonnes (DM)/ha/year of Spirodela from experiments that were carried out for
periods of 1-3 months in septage-fed 200 m2 ponds in Thailand; however, the yield
declined to the equivalent of ~9 tonnes (DM)/ha/year over a 6 months period. Based
on the available data and the foregoing discussion, it may therefore be concluded
that an average annual yield of around 10-20 tonnes/DM/ha can be obtained from an
aquatic environment where nutrients are generally not limiting and frequent harvesting
is practised to avoid plant overcrowding.

3.3.2 Duckweed farming


Duckweed farming is a continuous process requiring intensive management for
optimum production. Daily attention and frequent harvesting are needed throughout
the year to ensure optimum productivity. Duckweed can grow in water of any depth.

Table 3.5
Yields of various duckweed species under different environmental conditions
Species Environmental condition Yield (dry matter Reference
tonnes/ha/year)
L. minor UASB effluent 10.7 Vroon and Weller (1995)
L. minor Nutrient non-limiting water 16.1 Reddy and DeBusk (1984)
L. perpusilla Septage-fed pond 11.2 Edwards, Pacharaprakiti and
Yomjinda (1990)
L. perpusilla, S. Septage from septic tank 9.2-21.4 Edwards et al. (1992)
polyrrhiza and W.
arrhiza
Lemna Domestic wastewater 26.9 Zirschky and Reed (1988)
Lemna Sugar mill effluent 32.1 Ogburn and Ogburn (1994)
Lemna, Spirodela Domestic wastewater 13-38 Skillicorn, Spira and Journey
and Wolffia (1993)
Lemna and Wolffia Faecally polluted surface water 14-16 Edwards (1987)
S. polyrrhiza Domestic wastewater 17-32 Alaerts, Mahbubar and
Kelderman (1996)
S. polyrrhiza Sewage effluent 14.6 Sutton and Ornes (1975)
S. polyrrhiza Nutrient non-limiting water 11.3 Reddy and DeBusk (1985)
36 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

It will grow in as little as one centimetre of water. A pond depth of between 20


and 50 cm is generally recommended to reduce the potential sources of stress and to
facilitate harvesting (Gaigher and Short, 1986). Duckweeds are prone to be blown into
heaps by heavy winds or wave action. This allows light to penetrate the water column
and would stimulate phytoplankton and algal growth. If the plants become piled up in
deep layers, however, the lowest layer will be cut off from light and will eventually die
(Skillicorn, Spira and Journey, 1993). Plants pushed from the water onto a bank will
also dry out and die. Long narrow ponds that are sited perpendicular to the common
wind are recommended. Dividing the pond into smaller segments by using bamboo
can also mitigate the adverse effects of wind. An NGO called ‘PRISM’ applied a grid
of bamboo poles (Figure 3.4) of approximately 5 x 5 m in large ponds and wide canals.
This functioned satisfactorily for all conditions met up to that date in Bangladesh
(DWRP, 1998). Lemma USA Inc. promotes floating barrier grids made of polyethylene
that will reduce wind and wave action for its wastewater treatment plants. The sides
of the ponds must preferably be vertical to prevent the plants from becoming stranded
and at least 10 cm higher than the water level to accommodate heavy rains. The ponds
must be fed with effluent through furrows rather than pipes because the latter tend to
become clogged. Several inlets must be provided to spread the inflowing nutrients over
the pond.
Since the growth of duckweed is dependent on water temperature, pH and nutrient
concentration, these factors need to be balanced and maintained within reasonable
limits for duckweed to thrive. The management strategies for duckweed culture should
therefore focus on when to fertilize, harvest, and buffer; how much to fertilize and to
harvest; and which nutrients to supply. Appropriate management should be aimed at
maintaining a complete and dense cover of duckweed, low dissolved oxygen, and a
pH of 6-8. A dense cover shuts out light and suppresses the growth of algae, which
minimizes CO2 production from algal respiration and prevents its elevating effect on
pH.
Any waste organic material that is readily biodegradable and has a sufficiently high
nutrient content could be used for duckweed cultivation. The most economical sources
of such waste materials are all kinds of animal manure, kitchen wastes, wastes from a
wide range of food processing plants, biogas effluents, and slaughterhouse wastes. Solid
materials, such as manure from livestock, night soil from villages, or food processing
wastes, can also be mixed with water
and added to ponds at suitable levels.
Figure 3.4
A slowly flowing wastewater treatment canal covered All wastewater containing manure
with duckweed and provided with a bamboo base to or night soil must undergo an initial
prevent the duckweed from floating along the stream treatment by holding it for a few days
(PRISM Experimental Project, Mirzapur, Bangladesh) in an anaerobic pond, before using it to
cultivate duckweed.
Sutton and Ornes (1975) and Said et
al. (1979) demonstrated the necessity of
periodic additions of nutrients to small
duckweed culture systems receiving
municipal or dairy cattle wastes. Within
1-3 weeks, there was a noticeable drop
in N, P and K within the plants. There
was a corresponding drop in crude
protein as the plant nitrogen declined.
In unmanaged ponds, where duckweeds
are not routinely harvested, the plants
quickly become crowded and those
Source: DWRP (1998) beneath the surface die back.
Floating aquatic macrophytes – Duckweeds 37

Due to the high nitrogen requirement of duckweed and the relatively rapid loss
of nitrogen from aquatic system, this nutrient tends to be limiting in ponds fed with
wastewater (Gaigher and Short, 1986). Studies at Louisiana State University have
shown that the nitrogen conversion efficiency from agricultural waste to duckweed is
only about 30 percent under normal field conditions (Culley et al., 1981). Large-scale
duckweed production therefore requires the availability of relatively large quantities of
organic waste. The addition of cheap inorganic nitrogen could also therefore improve
the wastewater conversion efficiency. The other nutrients that are needed for optimum
growth of duckweed are phosphorus, potassium and trace minerals.

Fertilization
Urea is a suitable fertilizer, containing approximately 45 percent nitrogen, and is
rapidly converted to ammonia under normal conditions. Muriate of potash (MP) and
triple superphosphate (TSP) are commercial sources of potassium and phosphorus that
are widely available in most countries and have been used where duckweeds have been
farmed. Duckweed growth is not particularly sensitive to potassium or phosphorus
once an adequate threshold has been reached. A ratio of TSP to urea of 1:5 worked
satisfactorily in an experimental duckweed production programme in Bangladesh
(Skillicorn, Spira and Journey 1993). Similarly, a ratio for MP to urea of 1:5 was found
to be satisfactory for good production in the same duckweed production programme
in Bangladesh.
Nutrients are absorbed through all surfaces of the duckweed leaf (Leng, Stambolie
and Bell, 1995). There are at least three methods of fertilizer application including
broadcasting, dissolving in the water column of the plot, and spraying a fertilizer
solution on the duckweed mat.
A fertilizer application matrix aiming to achieve variable daily production ranging
from 500-1 000 kg of fresh duckweed per hectare was developed by PRISM in their
experimental programme at Mirzapur, Bangladesh (Table 3.6). Furthermore, PRISM
recommended daily fertilization rates for different types of duckweed (Table 3.7). The
application rate varies from 21-28 kg/ha/day (amounting to >7 tonnes/ha/year) with an
anticipated fresh biomass yield of 900-1 000 kg/ha/day. The daily fertilization rate for
duckweed cultivation developed by the Bangladesh Fisheries Research Institute (BFRI)
is presented in Table 3.8. The fertilizer schedules developed by PRISM and BFRI are
very similar (Tables 3.7 and 3.8), except that BFRI recommended half the dosage of
inorganic fertilizer when cow dung was used at the rate of 750 kg/ha/year.
Table 3.6
Dayly fertilizer application matrix for duck weed cultivation developed by PRISM in their
experimental programme at Mirzapur, Bangladesh
Fertilizer
application
500 600 700 800 900 1 000
(kg/ha)
Daily production of fresh plants (kg/ha)
Urea 10.0 12.0 14.0 16.0 18.0 20.0
TSP 2.0 2.4 2.8 3.2 3.6 4.0
MP 2.0 2.4 2.8 3.2 3.6 4.0
Sea salt 4.5 5.4 6.3 7.2 8.1 9.0

Source: Skillicorn, Spira and Journey, (1993)

Seeding
Seeding is a highly important management measure since a full duckweed cover should
be established before any algal bloom can start dominating the water body. The seed
rate advised is 60 kg/100 m2 for Spirodela spp. and Wolffia spp. and 40 kg/100 m2 for
Lemna spp. in order to obtain a dense cover in 3 days time (DWRP, 1998). From day
four onwards daily harvesting can start.
38 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

Stress management
Stress management of the crop is necessary particularly during very hot and dry
weather. ‘Dunking’ (dipping the duckweed below the water surface) once a day as
a regular crop maintenance practice is recommended; this reduces the stress from
overheating. Dunking consists of agitating the whole-cultivated area by hand until all
plants have been physically immersed and wetted.

Plant density and harvesting rate


The productivity of duckweed increases with increasing plant density up to a density
where the plants completely cover the surface of the water, and then remains constant.
In order to maintain good productivity and prevent competition by phytoplankton/
suspended algae, the density must be maintained at this level or a slightly higher level.
Competition between phytoplankton/suspended algae and duckweed is a potential
constraint to the cultivation of the latter in nutrient-rich water. Phytoplankton
smothers the roots of duckweed, which then turn yellow in colour, suffer a decline in
growth rate, and eventually die. The development of an algal bloom can also reduce
nutrient availability and thus eventually reduce the growth of duckweed.
An optimum standing crop density is a cover that is complete but which still
provides enough space to accommodate rapid growth of the colony. In the PRISM
experimental programme at Mirzapur, Bangladesh a base Spirodela density of 600 g/m2
was shown to yield a daily incremental growth of 50 to 150 g/m2/day (Skillicorn, Spira
and Journey, 1993). This is equivalent to a daily fresh (wet weight) crop production
rate of 0.5 to 1.5 tonnes/ha. These authors recommended a plant density of 400 to 800
g/m2 for optimum production. BFRI (1997) obtained duckweed production of 700-
1 500 kg/ha/day at plant densities varying from 400-600 g/m2 in their experimental
programme at Mymensingh.
High-density populations contain a high ratio of old fronds, which can be detrimental
in various ways. Duckweed should therefore be harvested frequently, preferably daily.
The standing crop density, or the weight of fresh plant per square meter, will determine
the amount and timing of harvests. Daily harvesting of the incremental growth of the
duckweed plot - averaging approximately 100 g/m2/day is recommended (Skillicorn,
Spira and Journey, 1993). Culley and Myers (1980) obtained an annual dry weight
production of 23.31 tonnes/ha with daily harvesting ranging from 10 to 35 percent of
the standing crop each day, depending on the season. Edwards (1990) recommended
25 percent harvesting of the duckweed biomass when duckweed growth completely
covers the pond, with the remaining 75 percent left in the pond for further growth.
Table 3.7
Rates of fertilization application for duckweed cultivation techniques developed by PRISM
Duckweed Rate of application (kg/ha/day)
Urea TSP MP
Spirodela 20 4 4
Wolffia 15 3 4
Lemna 15 3 3
Source: DWRP (1998)

Table 3.8
Rates of fertilization application for duckweed cultivation techniques developed by Bangladesh
Fisheries Research Institute (BFRI)
Fertilizer combination Rate of application (kg/ha/day)
Urea TSP MP Cow dung
Inorganic fertilizer only 15-20 3-4 3-4 -
Combination of organic and Inorganic fertilizer 7.5 1.5 1.5 750
Source: BFRI (1997)
Floating aquatic macrophytes – Duckweeds 39

This author opined that this harvest could be made every 1-3 days, depending on the
season.

Duckweed in wastewater treatment


Ferdoushi et al. (2008) tested the efficacy of Lemna and Azolla as biofilters of nitrogen
and phosphate in fish ponds in Bangladesh and found that they removed the excess
amount of nutrients from the water body and maintained sustainable environmental
conditions. Duckweeds have received much attention because of their potential to
remove contaminants from wastewater (Leng, Stambolie and Bell, 1995). Duckweed
wastewater treatment systems have been studied for dairy waste lagoons (Culley et al.,
1981), raw domestic sewage (Oron, 1994; Skillicorn, Spira and Journey, 1993; Alaerts,
Mahbubar and Kelderman, 1996), secondary effluent (Harvey and Fox, 1973), waste
stabilization ponds (Wolverton, 1979) and fish culture systems (Porath and Pollock,
1982; Rakocy and Allison, 1981). The basic concept of a duckweed wastewater
treatment system is to farm local duckweed on the wastewater requiring treatment.
Duckweed has a high mineral absorption capacity and can tolerate high organic loading
as well as high concentrations of micronutrients.
Duckweed wastewater treatment systems remove, by bioaccumulation, as much as
99 percent of the nutrients and dissolved solids contained in wastewater (Skillicorn,
Spira and Journey, 1993). These substances are then removed permanently from the
effluent stream following the harvesting of a proportion of the crop. The plants also
reduce suspended solids and BOD by reduction of sunlight in lagoons. Duckweed
systems distinguish themselves from other effluent wastewater treatment mechanisms
in that they also produce a valuable, protein-rich biomass as a by-product.
Depending on the wastewater, the harvested crop may serve as an animal feed, a
feed supplement supplying protein/energy and minerals, or a fertilizer. The question
of toxic elements must be considered if certain types of waste material serve as the
nutrient source for duckweed culture; for example, duckweed will absorb heavy metals
and insecticides from the wastewater. It may, therefore, have to be decontaminated
prior to feeding to animals if heavy metals are present in the water.
Landolt and Kandeler (1987) reported that of all aquatic plants, Lemnaceae have
the greatest capacity in assimilating the macro-elements N, P, K, Ca, Na and Mg.
Table 3.9 presents some data on daily nitrogen and phosphorus uptake efficiency by
duckweed. The results from the various studies are not comparable because different
species are used and different climatological and operational conditions were applied.
Temperature may have a significant effect on nutrient uptake efficiency as has also been
observed for other aquatic plants.
Table 3.9
Daily nitrogen and phosphorus uptake by duckweed
Region/Country Species Uptake (g/m2)
N P
Italy L. gibba/ L. minor 0.42 0.01
CSSR Duckweed 0.20 -
USA Lemna sp. 1.67 0.22
Louisiana, USA Duckweed 0.47 0.16
India Lemna sp. 0.50-0.59 0.14-0.30
Minnesota, USA Lemna sp. 0.27 0.04
Florida, USA S. polyrrhiza - 0.015
Source: adapted from DWRP (1997)

Culley et al. (1981) made a comprehensive study on nutrient uptake from wastewater
by a mixed culture of duckweed (Table 3.10). This shows that duckweeds are capable
of removing considerable amounts of organic wastes from natural water. An annual
40 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

nutrient removal capacity covered by Lemnaceae of 1 378 kg TKN, 345 kg P, and


441 kg K per hectare of water area was calculated.
Summarizing the results of PRISM Experimental Site at Mirzapur, Skillicorn, Spira
and Journey (1993) reported that treating an average flow of 125 m3/day of hospital,
school, and residential wastewater produced by a population of between 2 000 and
3 000 persons, the 0.6 ha duckweed treatment plant produces a final treated effluent
that exceeds the highest quality standards mandated in the USA (Table 3.11). These
authors also estimated that a typical duckweed wastewater treatment plant would yield
a daily harvest of up to one ton of duckweed plants (wet weight) per hectare or 90 kg
per hectare of dried, high protein duckweed meal each day.
Table 3.10
Mean annual dry duckweed yield and nutrient uptake by a mixed culture of duckweed1 harvested
daily2 from a 25 m2 lagoons3 receiving dairy cattle wastes4
Duckweed yield Duckweed5
(kg/m2) TKN (g/m2) P (g/m2) K (g/m2)
Dec-Feb 0.195 11.5 2.9 3.7
Mar-May 0.576 34.2 8.5 10.9
Jun-Aug 1.020 60.2 15.1 19.3
Sep-Nov 0.540 31.9 8.0 10.2
Total (kg/ha) 23 310 1 378 345 441
1
S. polyrrhiza, S. punctata, L. gibba and W. columbiana in approximate equal amounts at Baton Rouge, Louisiana,
USA (9-10 months growing season)
2
September-February: a mean of 105 of duckweed removed daily; March-August: a mean of 35 percent of
duckweed removed daily
3
Trials were run in triplicate
4
Fresh manure loading in first stage lagoon provided an effluent to the test lagoons with 15-65 mg/l TKN; 18-28
mg/l phosphorus (P); 38-69 potassium (K); pH 7.6-7.9
5
TKN 5.9 percent of dry weight, P 1.48 percent and K 1.89 percent
Source: modified from Culley et al. (1981)

Table 3.11
Quality of final treated effluent at Mirzapur Experimental Site on 23 March 1991
Treatment phase BOD5 NH3 (mg/l) P (mg/l) Turbidity
(mg/l) (FTU1)
Raw influent 120 39.40 1.90 113
Primary 60 32.20 2.00 85
Duckweed 1 0.03 0.03 10
US Summer Standards: Washington D.C. area 10 2.00 1.00 20
1
This turbidity unit standard is roughly equivalent to total suspended solids (TSS) times two
Source: Skillicorn, Spira and Journey (1993)

3.4 Chemical composition


Each frond of duckweed absorbs nutrients through the whole plant, not through a
central root system, directly assimilating organic molecules such as simple carbohydrates
and various amino acids. The entire body is composed of non-structural, metabolically
active tissue; most photosynthesis is devoted to the production of protein and nucleic
acids, making duckweeds very high in nutritional value. The nutritional content of
duckweed is probably more dependent on the mineral concentrations of the growth
medium than on the species or their geographic location. Water low in nutrients
generally results in reduced nutritional content in duckweed. Crude fibre content is
generally lower (varying between 7-10 percent) for duckweed grown in nutrient-rich
water than that grown in nutrient-poor water (11-17 percent).
Compared with most plants, duckweed leaves have little fibre (5 percent in cultured
plants) as they do not need to support upright structures (Leng, Stambolie and Bell,
1995). Crude fibre content was generally lower, varying between 7-10 percent, for
duckweed grown in nutrient-rich water as opposed to 11-17 percent for duckweed
Floating aquatic macrophytes – Duckweeds 41

grown in nutrient-poor water. In general, the ash content ranges between 12-18 percent
(Leng, Stambolie and Bell, 1995).
Duckweeds are known to accumulate large amounts of minerals in their tissues.
Skillicorn, Spira and Journey (1993) reported that fibre and ash contents are higher
and protein content lower in duckweed colonies with slow growth. Duckweeds are
rich source of nitrogen, phosphorous, potassium and calcium (Guha, 1997). The
concentration of N and P in duckweed tissues depend on the amount of N and P in
the water, up to a threshold concentration that has not been clearly defined. Above this
threshold, there is little increase in the tissue. Culley et al. (1978) suggested that under
lagoon conditions, 20-30 mg/l TKN might be required to maintain a crude protein
level above 30 percent. The crude protein content of duckweeds grown on various
nutrient solutions ranges from 7 to 45 percent of the plant dry weight, depending on
the nitrogen availability (Culley et al., 1981). When conditions are good, duckweed
contains considerable protein, fat, starch and minerals, which appear to be mobilized
for biomass growth when nutrient concentrations fall below the critical levels for
growth. Nutrient contents in duckweed may therefore vary according to the conditions
in which it is grown. Slow growth, starvation and aging have been reported to result in
protein levels as low as 7 percent DM (Landolt and Kandeler, 1987).
A summary of the nutritional composition of different species grown under different
environmental conditions is presented in Table 3.12. Fresh duckweed contained about
91-95 percent water and the moisture content is apparently not influenced by the
medium under which it was grown. Duckweed species grown under nutrient-poor
water or under sub-optimum nutrient conditions have crude protein contents varying
between 9-20 percent, while the level varied from 24-41 percent for duckweed species
grown in nutrient-rich water. The crude protein content of duckweed seems to increase
from trace ammonia concentrations to 7-12 mg N/L when crude protein reaches a
maximum of about 40 percent (Leng, Stambolie and Bell, 1995). Similarly, the lipid
content was lower (1.8-2.5 percent) in duckweed species grown in nutrient-poor water,
while it generally varied between 3-7 percent for duckweed grown in nutrient-rich
water. The medium in which duckweed was grown or the nutrient status of water did
not influence the ash content of duckweed (Leng, Stambolie and Bell, 1995). Skillicorn,
Spira and Journey (1993) reported that fibre and ash contents are higher and protein
content lower in duckweed colonies with slow growth.
Studies by Porath, Hepher and Koton (1979) and Rusoff, Blakeney and Culley
(1980) show clearly that the duckweed indeed has high quality protein. It has a better
essential amino acid profile than most plant proteins and more closely resembles
animal protein than any other plant proteins. According to Guha (1997), the protein
of duckweeds is rich in certain amino acids that are often rather low in plant proteins.
The nutritional value of Lemnaceae can be compared favourably with that of alfalfa in
terms of lysine and arginine, two amino acids important in animal feeds. Duckweeds
are rich in leucine, threonine, valine, isoleucine and phenylalanine and are low in
methionine and tyrosine.1 Some information on the amino acid content of various
aquatic macrophytes is contained in Annex 1. Annex 1 Table 3 shows mean values
determined for amino acids in four species of duckweed. It is evident that the values
for the essential amino acids compare favourably with the FAO reference pattern, with
the exception of methionine. The levels of amino acids are very similar in the various
species and all the essential amino acids were generally present.
Cultured duckweed has high concentrations of trace minerals and pigments,
especially β-carotene and xanthophyll (Haustein et al., 1988). Duckweeds store varying
amounts of calcium as calcium oxalate crystals in the vacuoles. Calcium oxalate may
be toxic in large doses and the amount should be reduced to make duckweeds more

1
www.mobot.org/jwcross/duckweed/nutritional-composition.htm
Table 3.12
42

Chemical analyses of various duckweed species grown under different environmental conditions
Duckweed species Aquatic environment Moisture Proximate composition1 (percent DM) Minerals1 Reference
(percent) (percent DM)
CP EE Ash CF NFE Ca P
L. gibba, USA Low nutrient lagoon2 n.s. 9.4 1.8 16.8 17.0 55.56 1.38 0.72 Culley et al. (1981)
L. minor, Bangladesh Pond, nutrient status not specified 92.0 14.0 1.9 12.1 11.1 60.9 n.s. n.s. Zaher et al. (1995)
L. minor, Bangladesh Ditch, nutrient status not specified 93.8 20.3-23.5 n.s. n.s. n.s. n.s. n.s. n.s. Majid et al. (1992)
L. polyrrhiza, India, raw leaf meal Freshwater 32.5 18.6 1.5 2.5 11.0 66.46 n.s. n.s.
L. polyrrhiza, fermented, India … n.s. 11.4 1.0 n.s. 7.5 n.s. n.s. n.s.
S. polyrrhiza, USA Low nutrient lagoon2 n.s. 13.1 2.5 13.3 16.1 55.06 1.21 0.56 Culley et al. (1981)
S. polyrrhiza, Bangladesh Ditch, nutrient status not specified 95.0 17.3-28.4 n.s. n.s. n.s. n.s. n.s. n.s. Majid et al. (1992)
S. punctata, USA Low nutrient lagoon2 n.s. 10.6 2.3 14.1 11.3 61.76 0.98 0.61 Culley et al. (1981)
W. arrhiza, whole plant, Bangladesh Ditch, nutrient status not specified 91.2 14.9 n.s. n.s. n.s. n.s. n.s. n.s. Majid et al. (1992)
L. gibba, USA High nutrient lagoon3 n.s. 36.3 6.3 15.5 10.1 31.86 1.81 2.60 Culley et al. (1981)
L. gibba, USA Dairy cattle waste lagoon n.s. 38.5 3.0 16.4 9.4 32.76 1.00 1.60 Hillman and Culley
(1978)
L. minima, USA Source not specified n.s. 31.0 2.0 14.0 10.0 42.0 n.s. n.s. Shireman, Colle and
Rottmann (1977)
L. perpusilla, Thailand Septage-fed earthen pond 94.0-94.3 25.3-29.3 3.8-4.5 15.4-17.6 6.9-7.6 n.s. n.s. n.s. Hassan and Edwards
1992)
S. oligorrhiza, USA Dairy cattle waste lagoon n.s. 37.8 3.8 12.0 7.3 39.16 1.30 1.50 Hillman and Culley
(1978)
S. oligorrhiza, USA4 Treated wastewater effluent n.s. 32.7 6.3 20.3 13.5 27.26 1.49 1.15 Culley and Epps (1973)
4
S. oligorrhiza, USA Untreated septic tank influent n.s. 32.3 n.s. n.s. n.s. n.s. 1.29 1.17 Culley and Epps (1973)
S. oligorrhiza, USA4,5 Anaerobic swine waste lagoon n.s. 41.4 5.1 12.9 8.3 32.36 0.91 2.07 Culley and Epps (1973)
S. polyrrhiza, Thailand Septage-fed earthen pond 91.0 23.8 3.8 18.3 11.7 42.46 n.s. n.s. Hassan and Edwards
(1992)
S. polyrrhiza, USA High nutrient lagoon3 n.s. 39.7 5.3 12.8 9.3 32.96 1.28 2.10 Culley et al. (1981)
S. polyrrhiza, USA Dairy cattle waste lagoon n.s. 40.9 6.7 12.9 8.7 30.86 2.10 1.40 Hillman and Culley
(1978)
S. punctata, USA High nutrient lagoon3 n.s. 36.8 4.8 15.2 9.7 33.56 1.75 1.50 Culley et al. (1981)
1
CP = crude protein; EE = ether extract; CF = crude fibre; NFE = nitrogen free extract; Ca = calcium; P = phosphorus
2
Low nutrient lagoon contained less than 5 mg/l TKN
3
High nutrient lagoon contained more than 30 mg/l TKN
4
Proximate composition and mineral content values corrected to dry basis
5
Mean of eight values sampled over five months period
6
Adjusted or calculated; not as cited in original publication
Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review
Floating aquatic macrophytes – Duckweeds 43

nutritious and digestible (Franceschi, 1989). The metabolic precursor of oxalate is L-


ascorbic acid (vitamin C). A study with water lettuce (Pistia stratiotes) (see section 5)
indicates that L-ascorbate and oxalate are synthesized within the crystal idioblast cells
(Kostman et al., 2001).

3.5 Use as aquafeed


Because of its attractive nutritional qualities and the relative Figure 3.5
Duckweed collected from
ease of production, a significant number of studies have been
the pond to be fed to tilapia,
carried on the potential utilization of duckweed biomass as Kumah Farms Complex
fish feed (Shireman, Colle and Rottmann 1977, 1978; Hillman (Kumasi, Ghana)
and Culley, 1978; Stephensen et al., 1980; Gaigher, Porath
and Granoth, 1984; Naskar et al., 1986; Hassan and Edwards,
1992). Available literature indicates that duckweeds are fed to
fish in fresh form as a sole feed or in combination with other
feed ingredients (Figure 3.5). Duckweeds are also fed as a dried
meal ingredient in pelleted diets. Intensive fish production
with duckweed as a predominant feed constituent has been
reported by a number of authors (Hepher and Pruginin, 1979;
Robinette, Brunson and Day, 1980; Culley et al., 1981; Landolt
and Kandeler, 1987, Skillicorn, Spira and Journey, 1993).
Research studies on the use of duckweed as fish feed have been
carried out under laboratory as well as under field conditions.
Successful results have also been obtained on the on-farm
utilization of duckweed as fish.

3.5.1 Laboratory studies


Successful feeding trials for grass carp with duckweed have been carried out since
the early 1960s. Studies on the consumption of duckweed by aquatic animals have
generally been confined to this species, although more recently feeding trials have also
been carried out with others, including common carp, catfish, Indian major carps and
tilapia.
Results on the use of duckweed as a feed for grass carp are generally very positive
(Galkina, Abdullaev and Zacharova, 1965; Nikolskij and Verigin, 1966; Fischer,
1968, 1970; Edwards, 1974; Porath and Koton, 1977; Shireman, Colle and Rottmann,
1977, 1978; Baur and Buck, 1980; Hajra and Tripathi, 1985). Galkina Abdullaev and
Zacharova (1965) reported that the grass carp showed more rapid growth when using
duckweed than other feed materials. Porath and Koton (1977) noted that the weight of
grass carp could be tripled (from 100 g to 300 g) within 50 days when feeding a mixture
of L. gibba and L. minor.
Fresh duckweeds have also been efficiently utilized by common carp, catfish, Indian
major carps and tilapia (Hepher and Pruginin, 1979; Robinette, Brunson and Day,
1980; Stephensen et al., 1980; Gaigher, Porath and Granoth, 1984; Naskar et al., 1986;
Hassan and Edwards, 1992).
Summary results of selected growth studies carried out on the use of fresh and dried
duckweed as feed for different fish species are presented in Tables 3.13 and 3.14. Fresh
and dried duckweed were fed to grass carp, Nile tilapia, common carp, Indian major
carps (rohu and mrigal), silver carp, Java barb, hybrid grass carp and hybrid tilapia.
The duckweed species evaluated were L. gibba, L. perpusilla, L. minima, L. minor,
Wolffia columbiana and W. arrhiza. Fresh duckweeds were fed as a sole feed whereas
dried duckweed meal was incorporated by partially replacing other conventional feed
ingredients in pelleted diets. Feeding trials were conducted for varying periods, ranging
from 60 to 155 days. Fish were fed ad libitum or at restricted level. In some studies,
44

Table 3.13
Performance of fish fed fresh duckweed
Duckweed/ Fish species Rearing system Rearing Control diet Feeding rate (percent Fish SGR SGR as FCR (DM Reference
days BW/day) size (percent) percent basis2)
Fresh Dry (g) of
weight weight control

L. gibba/ tilapia hybrid1 Recirculatory tanks 89 >1.0 2.66 0.67 1.0 Gaigher, Porath and
Granoth (1984)
L. gibba/ hybrid grass carp Static water outdoor 60 Catfish pellet Ad libitum 1 015 0.21 56 6.69 Cassani, Caton and
concrete tank (32 percent Hansen (1982)
protein)
L. perpusilla/ Nile tilapia Static water outdoor 70 2.5 26.3 0.97 2.2 Hassan and Edwards
concrete tank 5.0 27.4 1.09 3.7 (1992)
L. perpusilla/ Nile tilapia Static water outdoor 70 3.0 43.7 1.34 1.6 Hassan and Edwards
concrete tank 4.0 40.4 1.40 2.3 (1992)
L. minima/grass carp Flow-through circular 88 Ad libitum 2.7 3.74 1.72-1.97 Shireman, Colle and
fibre glass tank Rottmann (1977)
L. minima/grass carp Flow-through circular 68 Catfish chow Ad libitum 2.8 3.88 212 1.6 Shireman, Colle and
fibre glass tank (32 percent 62.8 1.19 212 2.7 Rottmann. (1978)
protein)
S. polyrrhiza/ Nile tilapia Static water outdoor 70 2.5 25.6 0.59 3.1 Hassan and Edwards
concrete tank 5.0 27.9 0.63 5.9 (1992)
W. columbiana/ hybrid grass Static water outdoor 60 Catfish pellet Ad libitum 1 033 0.51 135 3.76 Cassani, Caton and
carp concrete tank (32 percent Hansen (1982)
protein)
W. arrhiza/ grass carp Cement cistern 133 Ad libitum 5.5 3.50 Naskar et al. (1986)
W. arrhiza/ silver carp Cement cistern 133 Ad libitum 15.5 2.33 Naskar et al. (1986)
W. arrhiza/ common carp Cement cistern 155 Ad libitum 15.0 1.90 Naskar et al. (1986)
W. arrhiza/ Java barb Cement cistern 120 Ad libitum 9.5 2.49 Naskar et al. (1986)
W. arrhiza/ rohu Cement cistern 155 Ad libitum 5.0 2.87 Naskar et al. (1986)
W. arrhiza/ mrigal Cement cistern 155 Ad libitum 6.0 2.54 Naskar et al. (1986)
1
O. niloticus X O. aureus
2
Dry matter basis
Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review
Table 3.14
Performance of fish fed pelleted feeds containing dried duckweed
Duckweed/ Rearing Rearing Control diet Composition of test diet Inclusion Fish size SGR SGR as FCR Reference
Fish species system days level (g) (percent) percent of
(percent) control
L. minor/ Static 70 Fishmeal: sesame Duckweed meal incorporated by partial 13.5 2.55 2.13 98 2.1 Zaher et al.
Nile tilapia water oil cake: black gram replacement of black gram and rice (1995)
glass bran: rice bran bran
aquaria (27:25:19:25)
L. minor/ Static 140 Rice bran: groundnut Duckweed meal, groundnut cake, rice 40 3.0 2.29 94 3.1 Devaraj,
Common water cake (60:40) (17.5 bran and ragi flour (40:20:20:20) (21.3 Krishna and
carp outdoor percent protein) percent protein) Keshavappa
cement (1981)
cistern
Floating aquatic macrophytes – Duckweeds
45
46 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

the performances of fish fed duckweed were compared with control diets, although in
many of these studies no control diet was used for comparison.
Growth responses of different fish species fed various species of fresh duckweed
were variable. However, the general trend was that the grass carp performed better
than Nile tilapia and other species and the performances of duckweed as whole feed
were better than control diet (Table 3.13). Similarly, duckweed meal incorporated in
pelleted diets at 13.5 and 40 percent showed similar growth responses compared to the
growth responses of fish fed control diets (Table 3.14). The SGRs obtained for grass
carp fed fresh duckweed as whole feed varied between 1.2 and 3.9 while the SGR values
for Nile tilapia were between 0.6 and 1.4. Fasakin, Balogum and Fasuru (1999) reported
that duckweed meal (Spirodela polyrrhiza) can form up to 30 percent of the total diet of
Nile tilapia without significant effect on performance, compared to a control without
duckweed. However, inclusion levels above this level progressively decreased fish
performance.
Duckweed are generally the preferred macrophytes for most of the herbivorous fish,
although several authors reported that submerged macrophytes such as oxygen weed
(Hydrilla) and water velvet (Najas) are more preferred than others. The preference
of duckweed to other aquatic plants has been reported for grass carp and other fish
species in several observations (Opuszynsky, 1972; Duthu and Kilgen, 1975; Rifai,
1979; Cassani, 1981; Cassani and Caton, 1983). Information on whether fish prefer any
particular duckweed species over others is lacking.
Ad libitum feeding of fresh duckweed is mostly used for herbivorous fish. Limited
numbers of investigations have been carried out to optimize the feeding or consumption
rate of duckweed but most were carried out for grass carp and Nile tilapia. Nikolskij
and Verigin (1966) reported grass carp consumed fresh duckweed equal to their body
weight over a 24 hour period. Baur and Buck (1980) reported that grass carp consumed
from 85 percent to 238 percent of their body weight/day (BW/day) on a mixed diet
of Lemna, Spirodela and Wolffia spp. Shireman, Colle and Rottmann (1977) recorded
consumption rates varying from 7.2-7.4 percent BW/day on a dry weight basis (DW)
for grass carp while fresh duckweed (L. minima) was fed ad libitum. Since duckweed
contains about 92 percent moisture, the dry weight feeding rates given above are
equivalent to about 90-92 percent BW/day on a fresh weight basis. Shireman, Colle
and Rottmann (1978) fed fresh L. minima ad libitum to grass carp and recorded daily
mean consumption rates of 7.6 percent and 4.3 percent BW/day DW for 2.8 and
62.8 g sized fish respectively. Similar size-dependent feeding rates were reported by
Hassan and Edwards (1992) for Nile tilapia. These authors studied the effect of feeding
rate of L. perpusilla on the survival, growth and food conversion rate of Nile tilapia
and recorded that the optimal daily feeding rates of Lemna were 5, 4 and 3 percent
BW/day DW for fish of 25 to 44 g, 45 to 74 g and 75 to 100 g, respectively. Hassan
and Edwards (1992) concluded that duckweed should be fed to tilapia according to its
consumption rate, in order to avoid creating adverse water conditions, and that the
feeding rate should be decreased as the fish grow larger.
A study by Effiong, Sanni and Sogbesan (2009) also indicated that the inclusion of
duckweed meal in fish feeds could improve its binding potential and water stability.

3.5.2 Field studies and on-farm utilization


Several field studies and reports about the on-farm utilization of duckweed as feed
for various fish species exist (e.g. Edwards, 1980, 1987; Edwards, Pacharaprakiti and
Yomjinda, 1990; Skillicorn, Spira and Journey 1993; DWRP, 1998).
Edwards (1987) reported the on-farm utilization of duckweed in China and Taiwan
Province of China. This author reported that the duckweeds L. minor, S. polyrrhiza
and W. arrhiza are cultivated in small shallow ponds (similar to that illustrated in
Figure 3.6) fertilized with manure (livestock or human) and fed to grass carp fry
Floating aquatic macrophytes – Duckweeds 47

and fingerlings in nursery areas. Figure 3.6


Initially the fry are fed the smaller Two boys collecting duckweed from a village pond
(Jessore, Bangladesh)
Wolffia, but when they reach
6 to 7 cm in length they are fed
the larger Lemna and Spirodela.
In Taiwan Province of China a
mixture of Lemna and Wolffia
is cultivated in shallow earthen
ponds fed with faecally-polluted
surface water for use as fish feed.
Skillicorn, Spira and Journey
(1993) described the ‘duckweed-fed
carp polyculture system’ developed
in the PRISM experimental farm
at Mirzapur, Bangladesh. The
duckweed (Lemna, Spirodela and
Wolffia) carp polyculture model
has an 18-month cycle. Fingerlings
were introduced in August and
September and harvesting began in March and continued for approximately one year.
A second 18-month cycle began the following year and continued concurrently for
six months. After the initial six months, the model allowed year-round harvesting. Bi-
weekly harvesting was the preferred pattern, following a simple protocol to take the
largest fish (75 to 100 percentile) and the smallest (0 to 25 percentile) in each species.
The rationale was the assumption that the largest fish will exhibit a declining growth
rate and that the small fish are simply poor performers.
The production rates achieved in this programme suggested that one hectare of
duckweed production can support two hectares of carp polyculture. Empirical results
suggested that a polyculture stocked at about 30 000 fish/ha may be fed as much
duckweed as they will eat daily, regardless of the season. Fish were fed duckweed
throughout the day. Freshly harvested duckweed was brought in baskets to the pond
and distributed evenly among several ‘feeding rings or squares’ (Figure 3.7) consisting of
4 m2 open-bottom enclosures. Feeding rings provide access by the fish to the duckweed
and prevent it from dispersing over the pond surface. The feeding ring can be a
floating enclosure anchored
near the shore. Six feeding Figure 3.7
rings/ha were installed in Duckweed cultivated in an undrainable pond (Mymensingh,
Bangladesh)
the Mirzapur experimental
site and appeared to provide
sufficient access to food
for all fish. Figures 3.8 and
3.9 show the transport and
utilization of duckweed in
fish culture.
In the Mirzapur
experimental ponds, grass
carp was the primary
consumer of duckweed in
the polyculture. However,
both catla and common carp
also competed aggressively
for available duckweed feed
and consumed it directly.
48 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

Figure 3.8 Top-feeders directly


A duckweed collector carrying a bag of duckweed in a rickshaw van. These absorb about 50
professional duckweed collectors collect duckweed from various derelict
ponds and sell them to fish farmers (Jessore, Bangladesh)
percent of duckweed
nutrients in their
digestive systems.
Their faeces contain the
balance of the original
duckweed nutrients
and furnish relatively
high quantity detritus
to the bottom-feeders.
A duckweed-fed fish
pond thus appears to
provide a complete,
balanced diet for those
carp that consume
it directly, while the
faeces of duckweed-
feeding species, which
are consumed directly
by detritus feeders or
indirectly through fertilization of plankton and other natural food organisms,
provide adequate food for the remaining bottom and mid-feeding carp varieties. The
fertilization of a duckweed-fed fish culture is therefore indirect and gradual, resulting
from bacterial decomposition of fish faeces, dead algae, and other fermenting organic
material.
Skillicorn, Spira and Journey (1993) reported that
Figure 3.9 the first annual cycle of carp production produced
A farmer is releasing a bag of duckweed
slightly more than 10 tonnes/ha/year. However, these
to his carp pond. These duckweeds are
generally purchased from a group of authors opined that a yield of between 10 to 15 tonnes/
professional duckweed collectors ha/year appears to be sustainable before biological
(Jessore, Bangladesh) constraints become limiting factors.
DWRP (1998) reported further follow-up of the
duckweed-fed carp polyculture system developed by
PRISM in Bangladesh. This report included the results
of the demonstration farms as well as the results of the
farmers’ ponds. The duckweed-fed carp polyculture
system practised by PRISM had two distinct differences
from the model described by Skillicorn, Spira and
Journey (1993). Apparently, PRISM included Nile
tilapia with the traditional six-species carp culture
system and provided other supplemental feed along with
the duckweed. The farmers that adopted duckweed-
based aquaculture produced an average of 3.6 tonnes/
ha/year in comparison to the national average fish
production of 2.1 tonnes/ha/year in Bangladesh at that
time. PRISM itself achieved a production level of 11
tonnes/ha/year in 1993 and 16 tonnes/ha/year in 1996
in its demonstration farm.
The results of the duckweed-based carp polyculture
of PRISM are presented in Tables 3.16, 3.17 and 3.18.
Table 3.15 presents fish stocking and harvesting data in
1994. Tilapia were not stocked but multiplied on their
Floating aquatic macrophytes – Duckweeds 49

own. Data for feed application, fish yield and food conversion ratio in farmers’ ponds
and in demonstration farms of PRISM are presented in Tables 3.16 and 3.17. The specific
influence of duckweed feeding on yield and food conversion was obscured because
other supplemental food was also added to the fish ponds. In neither demonstration
ponds nor farmers’ ponds could any example be found of pure duckweed feeding.
Table 3.17 presents data collected from demonstration farm production in Mirzapur
over a period of three years. The results from fish ponds fed duckweed from organic
wastewater plants have been kept separate from those fed on duckweed grown on
chemical fertilizers. Whether there has been any difference in nutritive value between
the duckweed from these different treatments could not be checked. What is interesting
is to compare the difference in the ratio of duckweed to the other supplemental feed
that was being applied. At first the information in Table 3.17 suggests that higher
proportions of duckweed influence the conversion rate adversely. This conclusion
may not be true, however, since the total amount of food applied in the ponds treated
with waste-grown duckweed was clearly too high. Since the ponds did not show any
increased production with a high rate of feeding, it must be assumed that they were
at their carrying capacity most of the time and that all the extra food offered was
apparently wasted. This seems to imply that the sustainable level of fish production
from a duckweed-based polyculture lies around 10-15 tonnes/ha/year.
Table 3.15
The ratio of fish species stocked and harvested by PRISM in 1994
Species Stocking rate (percent) Harvest rate (percent)
Tilapia 0 38.8
Catla 20 6.7
Rohu 20 9.7
Mrigal 20 9.3
Silver carp 15 24.3
Grass carp 20 7.3
Common carp 5 3.3
Other 0 0.6

Table 3.16
Feed application, fish yield and food conversion ratio in farmers’ ponds in two locations in
Bangladesh during 1995-96
Location Oil cake Wheat bran Duckweed Total Fish yield FCR
(kg/ha DM) (kg/ha DM) (kg/ha DM) (kg/ha DM) (kg/ha)
Tangail 2 742 1 441 1 526 5 833 3 290 2.1
Manikganj 2 556 1 854 1 465 5 874 5 007 1.5
Source: DWRP (1998)

Table 3.17
Feed application, fish yield and food conversion ratio in demonstration ponds at Mirzapur
Experimental Site, Bangladesh during 1993-95
Nutrient Season Oilcake Wheat Duckweed Total percent Fish FCR
source of (kg/ha bran (kg/ha DM) (kg/ha duckweed yield
duckweed DM) (kg/ha DM) used in (kg/ha)
DM) feed
Chemical 1993 8 504 1 065 6 662 16 231 41 13 430 1.2
1994 11 722 507 5 902 18 131 33 15 080 1.2
1995 12 107 122 5 810 18 039 32 11 520 1.6
Wastewater 1994 9 810 - 19 840 29 650 67 10 580 2.8
1995 18 307 - 23 300 41 607 56 12 620 3.3
Source: DWRP (1998)

The food conversion values obtained when various duckweed species were fed to
different fish species are presented in Table 3.18. Duckweeds were fed mostly in the
fresh form and most of the values available are for grass carp and Nile tilapia. The
50 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

values are variable, but the available data does not indicate if the variability was due to
the fish species or to the duckweed species used. Generally, most FCRs are between 2.0
and 3.0, although an FCR of 1.0 was reported for hybrid tilapia and a very high FCR
(6.7) for hybrid grass carp when both were fed L. gibba. This latter result was probably
due to size of the fish (>1.0 kg) used in the feeding trial. Shireman, Colle and Rottmann
(1978) reported an FCR of 1.6 for 2.8 g grass carp when fed fresh L. minima but the
FCR was 2.7 for 63 g fish. Hassan and Edwards (1992) reported that food conversion
was significantly affected by the feeding rate. For example, the FCR was 3.1 when
S. polyrrhiza was fed to Nile tilapia at a feeding rate of 2.5 percent BW/day whereas
it was 5.9 at a feeding rate of 5.0 percent BW/day. Similarly, FCR increased from 2.2
to 3.7 with an increase in feeding rate from 2.5 to 5.0 percent BW/day for Nile tilapia
when fed L. perpusilla.
Generally, the FCR values reported for duckweed-based polyculture of carps in
Bangladesh (Tables 3.16 and 3.17) were very good, being between 1.5 and 2.1 for
farmer’s ponds and 1.2 to 1.6 for demonstration ponds. However, it must be pointed
out that duckweed was not used as the sole feed in these ponds, which were usually
fertilized in addition to the use of oilcake, rice bran and wheat bran as supplemental
feeds. Low FCRs for duckweed may be expected, since these plants have relatively low
fibre and high protein contents (Table 3.12) and a good amino acid profile (Annex 1
Table 3). Although it is difficult to generalize from the available data, an FCR value of
2.5 may be a reasonable expectation for grass carp and Nile tilapia based.
Table 3.18
Food conversion ratios of duckweed to fish
Duckweed Fish species Fish size Food conversion Reference
(g) ratio (FCR)
DM2 FW2
L. gibba, fresh Tilapia hybrid 2.7 1.0 Gaigher, Porath and
Granoth (1984)
L. gibba, fresh Hybrid grass 1 015 6.7 Cassani, Caton and
carp Hansen (1982)
L. perpusilla, fresh Nile tilapia 8-10 3.7 60.6 Edwards, Pacharaprakiti
and Yomjinda (1990)
L. perpusilla, fresh Nile tilapia 26-27 2.2-3.7 Hassan and Edwards
(1992)
L. perpusilla, fresh Nile tilapia 40-44 1.6-1.9 Hassan and Edwards
(1992)
L. minima, fresh Grass carp 2.7 1.7-2.0 Shireman, Colle and
Rottmann (1977)
L. minima, fresh Grass carp 2.8 1.6 Shireman, Colle and
Rottmann (1978)
L. minima, fresh Grass carp 63 2.7 Shireman, Colle and
Rottmann. (1978)
L. minor, fresh Nile tilapia n.s. 33 Rifai (1979)
L. minor, dried Nile tilapia 2.5 2.1 Zaher et al. (1995)
L. minor, dried Common carp 3.0 3.1 Devaraj, Krishna and
Keshavappa (1981)
Lemna sp., fresh Grass carp n.s. 37 Hepher and Pruginin
(1979)
S. polyrrhiza, fresh Nile tilapia 26-28 3.1-5.9 Hassan and Edwards
(1992)
S. polyrrhiza, dried, All-male 13.9 2.0 Fasakin, Balogun and
30 percent inclusion tilapia Fasuru (1999)
W. arrhiza, fresh Six carp 5.0-15.5 5.6 78.8 Naskar et al. (1986)
species1
W. columbiana, fresh Hybrid grass 1 033 3.8 Cassani, Caton and
carp Hansen (1982)
Mixture of Lemna, Grass carp n.s. 1.6-4.1 Baur and Buck (1980)
Spirodela and Wolffia
1
Polyculture of six carp species (grass carp, silver carp, common carp, Java barb, rohu and mrigal)
2
FW = fresh weight basis; DM = dry matter basis
Floating aquatic macrophytes – Duckweeds 51

Digestibility coefficients of Lemna, Spirodela and Wolffia fed to grass carp, tilapia
and rohu are presented in Table 3.19. Considering the importance of duckweed as fish
feed, it is surprising to note that only a few studies have been carried out to investigate
its digestibility for fish. Van Dyke and Sutton (1977) were probably the first to
investigate the digestibility of duckweed (mixture of L. minor and L. gibba) in detail
for grass carp. These authors estimated the true dry matter digestibility of duckweed to
be 65 percent, while the apparent digestibility was 53 percent for dry matter, 80 percent
for crude protein, 58 percent for organic matter, 26 percent for ash and 61 percent for
gross energy. The dry matter digestibility of L. gibba, S. polyrrhiza and W. arrhiza for
grass carp found by other authors (Table 3.19) varied between 67-82 percent, while
the dry matter digestibility of L. gibba for hybrid tilapia was reported to be
65 percent. Grass carp passes its food rapidly through a short, unspecialized gut
and the fish probably does not produce cellulase (Van Dyke and Sutton, 1977); it is
therefore unrealistic to expect that more than 50-60 percent of the feed consumed
would actually be digested.
Table 3.19
Digestibility of duckweed for selected fish species
Fish Fish size Digestibility
Duckweed species (g) (percent)1 Reference
DM CP EE NFE CF GE
L. gibba Hybrid 2.7 65 86 Gaigher, Porath and
tilapia2 Granoth (1984)
L. gibba Grass 82 Lin and Chen (1983,
carp cited by Wee, 1991)
L. gibba and Grass 320 53 80 61 Van Dyke and Sutton
L. minor (1:1) carp (1977)
S. polyrrhiza Grass 75 Lin and Chen (1983,
carp cited by Wee, 1991)
W. arrhiza Grass 67 Lin and Chen (1983,
carp cited by Wee, 1991)
W. arrhiza Rohu 3.6 91.5 93.5 81.2 84.4 Ray and Das (1994)
1
DM = dry matter; CP = crude protein; EE = ether extract; CF = crude fibre; NFE = nitrogen free extract; GE = gross energy
2
O. niloticus X O. aureus
53

4. Floating aquatic macrophytes


– Water hyacinths

Mature plants of water hyacinths Figure 4.1


Part of River Yamuna covered with lush green water
(Eichhornia crassipes) consist of long,
hyacinth, Delhi, India
pendant roots, rhizomes, stolons, leaves,
inflorescences and fruit clusters. The
plants may be up to 1 m high, although
40 cm is the more usual height. The
inflorescence bears 6-10 lily-like flowers,
each 4-7 cm in diameter. The stems and
leaves contain air-filled tissue, which
gives the plant considerable buoyancy.
Vegatative reproduction takes place at a
rapid rate under preferential conditions
(Herfjord, Osthagen and Saelthun,
1994).
Water hyacinths are considered as
nuisance species because they multiply
rapidly and clog lakes, rivers and ponds. Courtesy of Rina Chakrabarti

The thick mats (Figure 4.1) formed


under favourable conditions often obstruct fishing, shipping and irrigation and are
hard to eradicate. Great efforts are being made to contain water hyacinths but, on
the other hand, attempts are being made to find practical uses for the large biomass
that is available. It offers the potential for use as fodder for domestic animals, as fish
feed, for the production of biogas and for the removal of heavy metals and phenols
from polluted waters. For example, studies have shown that about 1 million L/day
of domestic sewage could be treated over an area of 1 ha through water hyacinths,
reducing the BOD and COD by 89 and 71 percent, respectively (Reddy et al., 2005).

4.1 Classification
There are seven species of water Figure 4.2
hyacinth, the best known being the Common water hyacinth (Eichhornia crassipes)
common water hyacinth, Eichhornia
crassipes, which is a perennial free-
floating aquatic plant belonging to the
family Pontederiaceae (Figure 4.2).

4.2 Characteristics
4.2.1 Importance
Water hyacinths are found in most of
the tropical and subtropical countries
of the world. According to Mitchell

Source: USDA
54 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

(1976), the water hyacinth is indigenous to South America, particularly to the


Amazonian basin. It started its worldwide journey as an ornamental plant when first
introduced into the USA in 1884 (Penfound and East, 1948 cited by Edwards, 1980). It
reached Australia in 1895, India in 1902, Malaysia in 1910, Zimbabwe in 1937 and the
Republic of the Congo in 1952.

4.2.2 Environmental requirements


According to Wilson et al. (2001) there are five main factors limiting the growth rate
and carrying capacity of water hyacinth: salinity, temperature, nutrients, disturbance
and natural enemies.
Water hyacinths seem unable to survive salinities above 2 ppt. Olivares and
Colonnello (2000) reported that water hyacinth survives salinities of 1.3-1.9 ppt in the
Orinoco (South America) and Kola (1988) reported that the plant grew well at salinities
below 1 ppt.
Low temperatures stop the plant establishing in temperate areas and prevent it from
reaching high levels in the sub-tropics. Knipling, West and Haller (1970) proposed
a parabolic relationship between temperature and growth rate, with growth tailing
off quickly after the optimum of 30 ºC. Imaoka and Teranishi (1988) proposed that
the intrinsic growth rate, r, increases exponentially with ambient temperatures in the
range 14-29 ºC, growth ceasing below 13 ºC. Frost is a major cause of leaf mortality
in temperate regions. Applying mathematical modelling, using existing data, Wilson,
Holst and Rees (2005) examined the role of two important environmental factors,
temperature and nutrient level, on the growth of water hyacinths. Their model
predicted a linear reduction in specific growth rate with density. These authors set the
minimum (Qmin), optimum (Qopt) and maximum (Qmax) temperatures for water hyacinth
as 8, 30 and 40 ºC, respectively. The growth of water hyacinths is affected by low air
humidity, ranging from 15-40 percent relative humidity (Freidel and Bashir, 1979).
The levels of available nitrogen and phosphorous are the most important factors
limiting growth (Wilson et al. 2001). The half-saturation co-efficients for water
hyacinths grown under constant conditions have been found to be from 0.05-1 mg/l
for total nitrogen and from 0.02-0.1 mg/l for phosphates. Growth quickly tails off
below the lower limits. Wilson, Holst and Rees (2005) suggested that nitrogen is
limiting if total nitrogen concentration is less than seven times that of the phosphorus
concentration. Water hyacinths show logistic growth. The model assumed that plants
grow in the absence of interspecific competition. In fact, the plant soften grow in areas
previously free of aquatic vegetation.
Flooding can break up large mats of water hyacinth and leave plants stranded on
land (Wilson et al. 2001). Wave action may limit growth by directly damaging plants
and by forcing the weed to maintain aerenchymatous tissue.

4.3 Production
Water hyacinth grows in all types of freshwater, lentic and lotic. Westlake (1963)
predicted that water hyacinths might be exceptionally productive plants since they are
warm water species with submerged roots and aerial leaves like emergent macrophytes.
Production statistics of this macrophyte in various aquatic environments are available
(Table 4.1). The productivity varies widely and is dependent on the environment under
which it grows. Wolverton and McDonald (1976) reported a yield of water hyacinth of
up to 657 tonnes/ha/year DM in ponds fertilized with sewage nutrients, while Coche
(1983) reported an even higher yield of 750 tonnes/ha/year in irrigation canals in
China. However, many of these reported yields are extrapolated. It may therefore not
be possible to obtain the higher calculated productivities on a large scale, since it would
be difficult to maintain the most rapid growth rates obtained on a small experimental
scale throughout the year (Edwards, 1980). The latter author, however, opined that an
Floating aquatic macrophytes – Water hyacinths 55

annual production of 200 tonnes/ha/year might be attainable in eutrophic waters in


the tropics.
Table 4.1
Productivity of water hyacinths under different aquatic environments
Aquatic environment Yield (tonnes/ha/year)
Fertile ponds 15-200
Artificially fertilized ponds 75.6-191.1
Fertilized pond 70.8
Fertilized pond with sewage effluent 212-657
Fertilized pond with sewage effluent 219
Irrigation canals in China 400-750
Nutrient non-limiting water of Florida, USA 106
Man-made lakes of central Java 255
Source: Edwards (1980); Little and Muir (1987)

China is probably the only country where water hyacinth has been reported to be
cultivated with two other aquatic macrophytes, namely water lettuce (Pistia stratiotes)
and alligator weed Alternathera philoxeroides (Edwards, 1987). These plants are usually
cultivated in rivulets, small bays or swamps, and are usually fed to pigs.

4.4 Chemical composition


A summary of the chemical composition of water hyacinths (fresh, dried and
composted) from different geographic regions of the world is presented in Table
4.2. Like most other aquatic macrophytes, water hyacinths have very high moisture
content; the dry matter generally varies between 5-9 percent. Table 4.2 indicates that
there is little variation in proximate composition in relation to geographic location.
Variation, however, does exist between the proximate composition of whole plants
and leaves. The crude protein content of the whole plant is about 12-20 percent DM,
although a level as low as 9 percent was reported in studies. Gohl (1981) reported that
the crude protein of fresh green part of water hyacinths from India and the Philippines
was 12.8-13.1 percent DM. The crude protein content of leaf meal appears to be higher
than the whole plant and varies between 20-23 percent.
Like most other aquatic macrophytes, the crude lipid content of water hyacinths
is usually low and varies between 2-4 percent on dry matter basis regardless of whole
plant or leaves. The ash content of whole plants varies between 15-34 percent while
it is between 10-18 percent for leaves. Crude fibre content is usually high in water
hyacinths and ranges between 17-32 percent, irrespective of whole plant or leaves.
Some information on the amino acid content of various aquatic macrophytes is
contained in Annex 1.
Gunnarsson and Petersen (2007), in a review that covered water hyacinths collected
from various sources, also reported levels of some other components: hemicellulose
22-43.4 percent; cellulose 17.8-31 percent; lignin 7-26.36 percent; and magnesium 0.17
percent. Matai and Bagchi (1980) provided some additional component levels for fresh
water hyacinths, namely that the ash contained 28.7 percent K2O, 1.8 percent Na2O
and 21 percent Cl.

4.5 Use as aquafeed


Because of their relatively high protein content and abundance in tropical and sub-
tropical countries, a significant number of research studies have been carried out to
find the potential for the utilization of water hyacinths as a fertilizer, for example
by Sipauba-Tavares and Braga (2007) for the rearing of tambaqui (Colossoma
acropomum), and as a fish feed in pond aquaculture. Available literature indicates
that water hyacinths are fed to fish either in fresh form, or as a dried meal in pelleted
diets, or composted as feed and fertilizer. Apart from these three forms, attempts are
56

Table 4.2
Proximate composition and mineral content of water hyacinths
Form DM (%) Proximate composition1 (% Minerals (% DM) Reference
DM)
CP EE Ash CF NFE Ca P
Fresh whole plant, Vietnam 5.8 13.4 2.5 34.1 31.8 18.2 n.s. n.s. Tuan et al. (1994)
Dried whole plant, Thailand 89.6 15.8 3.9 30.3 22.8 27.23 n.s. n.s. Edwards, Kamal and Wee (1985)
Whole plant, sun-dried, Thailand 90.2 8.9 1.0 17.3 23.0 49.8 n.s. n.s. Klinavee, Tansakul and Promkuntong (1990)
Fresh leaf, Thailand 7.3 22.7 3.9 18.8 17.5 37.13 1.79 0.83 Somsueb (1995)
Leaf meal, Bangladesh2 89.3 29.7 3.6 10.2 17.2 39.3 1.15 0.64 Hasan (1990)
Leaf meal, India 91.8 22.8 0.8 13.1 19.4 43.9 n.s. n.s. Murthy and Devaraj (1990)
Leaf meal, India 90.8 19.7 2.8 10.2 20.0 47.33 n.s. n.s. Nandeesha et al. (1991)
Whole plant, composted, Thailand 89.5 15.8 1.4 49.8 10.4 22.63 n.s. n.s. Edwards, Kamal and Wee (1985)
Leaf protein concentrate, USA n.s. 32.1 7.7 7.3 7.9 45.0 n.s. n.s. Liang and Lovell (1971)
Fresh whole plants 4.5 9.4 n.s. 24.2 n.s. … 2.21 0.74 Matai and Bagchi (1980)
Fresh whole plants, various locations 6.2-9.4 11.9-20.0 3.5 15.0-25.7 18.9 … 0.58 0.53 Gunnarsson and Petersen (2007)
Mean from various sources 8.1 15.5 1.9 17.8 25.3 39.53 2.06 0.6 Hertrampf and Piedad-Pascual (2000)
1
CP = crude protein; EE = ether extract; CF = crude fibre; NFE = nitrogen free extract; Ca = calcium; P = phosphorus
2
Leaves collected from a particular type of water hyacinth with a long and thin stem
3
Adjusted or calculated; not as cited in original publication
Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review
Floating aquatic macrophytes – Water hyacinths 57

also made to feed water hyacinths to fish by processing them with other techniques.
Many of these studies were conducted under laboratory conditions and reports of on
farm utilization as fish feed are rather limited. Information on these topics has been
grouped into several sections: four dealing with the various forms of water hyacinth
(fresh, dried, composted and fermented, and other processing techniques), followed by
comments on food conversion efficiency and digestibility.

4.5.1 Fresh form


Many researchers have investigated the use of water hyacinth in its fresh form. The
high moisture content is a major constraint in its use as fish feed, which has proved to
be unsuccessful in many cases. Hyacinth leaves are generally cut into small pieces and
fed to grass carp or other macrophytophagous fish. Generally, grass carp feed on this
plant only when no other macrophytes or feeds are available.
Riechert and Trede (1977) reported the results of a preliminary indoor laboratory
trial carried out in Germany on the feeding of water hyacinths to grass carp. Eleven
month old fish weighing 38 to 104 g were fed for 50 days exclusively on water
hyacinths. Roots and leaves were accepted readily by the grass carp but the swollen
petioles reluctantly. The fish grew well, producing 6.5 g live weight from 10 g DM
hyacinth (FCR = 1.54). These authors also noted that grass carp above 80-100 g were
better able to utilize hyacinth leaves compared to smaller fish and postulated that only
50-60 percent of the feed consumed was actually digested.
Tuan et al. (1994) used both fresh and fermented water hyacinth as supplementary
feed in nursery ponds in Vietnam for fingerlings (1-6 g) of Nile tilapia, common carp,
grass carp and Java barb. Fresh whole water hyacinth was chopped and mixed with
rice bran at a ratio of 2:1 or 1:1 and fed to fish. A water hyacinth-rice bran mix was
also fermented and fed. The growth of fish obtained by feeding the hyacinth-rice bran
mixture was comparable to the growth obtained from rice bran alone. Rice bran is
normally applied to nursing ponds in Vietnam. In terms of weight gain and specific
growth rate, water hyacinths mixed with rice bran at a ratio of 2:1, either raw or
fermented, could be used to replace rice bran in nursery ponds. Amongst the four
species used, Nile tilapia performed better than the other species, exhibiting a specific
growth rate of 4.3-4.8 percent/day. The specific growth rates of grass carp, Java barb
and common carp were 4.06-4.19 percent, 2.84-3.00 percent and 2.49-2.66 percent per
day, respectively.
As noted above, the use of fresh water hyacinth as fish feed has achieved limited
success, principally because of its high moisture content. There are several other
limitations to its use for this purpose. For example, the fresh plant contains prickly
crystals, which make it unpalatable (Gohl, 1981). This was thought to be probably due
to the presence of raphids and oxalates in water hyacinths (Dey and Sarmah, 1982).
Microscopic examination of water hyacinths reveals the presence of sharp needles
formed by calcium oxalate, which may be harmful for fish (Bolenz, Omran and
Gierschner, 1990).
Fresh whole water hyacinth has been applied to ponds as feed and fertilizer in
China, but the fish were reluctant to accept it and it took a long time to decompose,
eventually resulting in inefficient utilization (Anonymous, 1980). Several processing
techniques have therefore been employed to increase its nutritive value and to decrease
the high moisture content. These include its use in dried and composted forms, and
the incorporation of leaf meal in pelleted feeds. Another practice prevalent in China is
the application of paste or mashed water hyacinth, which releases the mesophyll cells
in water for consumption by carps. The processing methods employed so far and the
results achieved with various fish species are summarized in subsequent sections.
58 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

4.5.2 Dried meal form


One of the most commonly used methods for processing of water hyacinth is drying.
In tropical and sub-tropical countries, water hyacinths are often sun-dried, as other
drying methods can be expensive. Two days of good sun drying would be sufficient
to reduce the moisture content to about 10-12 percent. A number of growth studies
have been conducted under laboratory conditions using dried water hyacinth in
pelleted feeds for carps, tilapia and catfish. In most cases the dried water hyacinth was
ground into a meal and fed to fish, partially or completely replacing fishmeal or other
conventional protein sources.
A summary of the results of the selected growth studies carried out on the use of
dried water hyacinth meal in pelleted feeds for different fish species is presented in
Table 4.3. Whole water hyacinth or its leaf meal was evaluated as a major ingredient
in pelleted diets for Nile tilapia (Oreochromis niloticus), Java tilapia (O. mossambicus),
grass carp (Ctenopharyngodon idella), common carp (Cyprinus carpio), the Indian
major carp rohu (Labeo rohita), stinging catfish (Heteropneustes fossilis), Java barb
(Barbonymus gonionotus), sepat rawa (Trichogaster sp.), matrincha (Brycon sp.) and
African catfish (Clarias gariepinus). The dietary incorporation level of water hyacinth
meal used varied widely, ranging from as low as 2.5 percent to as high as 100 percent.
In most of these studies, the performance of fish fed diets containing various inclusion
of water hyacinth was compared with the use of control diets. Various types of control
diets were used, including commercial pellets, fishmeal-based pellets, the traditionally
used rice bran-oil cake mixtures, and a mixture of fishmeal and cereal by-products.
Growth responses of different fish species fed test diets containing different
inclusions of water hyacinth meal have been highly variable. For example, significant
reduction in growth responses were reported by Hasan, Moniruzzaman and Omar
Farooque (1990) for rohu fry and by Hasan and Roy (1994) for rohu fingerling when
27-30 percent water hyacinth leaf meal was included to replace the fishmeal protein
of the control diet. Similarly, Klinavee, Tansakul and Promkuntung (1990) recorded
significant reduction in growth responses of Nile tilapia when fed a test diet containing
40 percent water hyacinth meal. However, Murthy and Devaraj (1990), using a 50
percent dietary inclusion level in diets for grass carp and common carp, Dey and Sarmah
(1982) using 100 percent inclusion for Java tilapia, and Saint-Paul, Werder and Teixeira
(1981), using 18.5 percent inclusion for matrincha (Brycon sp.), respectively recorded
either similar or higher growth responses compared to control diets. However, in some
of these studies, the control diet consisted only of a rice bran-oil cake mixture, which
may itself have not generated good growth. Edwards, Kamal and Wee (1985) tested the
growth response of Nile tilapia to 75 and 100 percent displacement of a 32.5 percent
protein commercial tilapia pellet by water hyacinth meal. The test diets resulted in
only a 10-15 percent reduction in SGR. This is an interesting performance for water
hyacinth meal. However, these authors concluded that although the experimental fish
obtained their nutrition directly from the diets, they must also have obtained some
indirect nutrition from the plankton in the static water experimental system used. This
assumption of indirect nutritional benefit from phytoplankton may also have been
true in the experimental studies conducted by Dey and Sarmah (1982) and Murthy and
Devaraj (1990).
Hertrampf and Piedad-Pascal (2000) suggested inclusion rates for water hyacinth
in farm-mixed feeds for the farming of herbivorous or omnivorous freshwater fish in
simple farming systems where it is available at low cost. These authors recommended
that suitable inclusion levels were either 25-50 percent as a supplementation of basic
feed (e.g. rice bran, broken rice, chicken manure) or 5-10 percent as a replacement
protein source in formulated feeds (fish meal, vegetable oil meals/cake).
Table 4.3
Performance of different fish species to pelleted feeds containing dried water hyacinth meal
Fish species Rearing system Rearing Control diet Composition of test diet Dietary WH Fish size SGR SGR as FCR Reference
days inclusion % of
(g) (%)
level (%) control
Nile tilapia Static water 84 Commercial pellet WH meal incorporated at two 75 14.44 1.80 90.5 2.41 Edwards, Kamal and Wee
outdoor concrete (32% protein) levels in control diet by simple 100 13.50 1.70 85.4 2.61 (1985)
tank displacement
Nile tilapia Clear water fibre 77 Chicken pellet (16.8 WH leaf meal, fishmeal and rice 40 7.70 1.01 64.0 4.3 Klinavee, Tansakul and
glass tank protein) bran (40:18.5:34.3) Promkuntong((1990)
Java tilapia Indoor static glass 28 Rice bran: mustard Dried WH petiole meal 100 15-22 -2 n.s n.s. Dey and Sarmah (1982)
aquaria oil cake (50:50)
Java tilapia Cages in lake 70 No control diet WH meal, fishmeal, corn meal and 2.5 10-20 1.34 n.s n.s Hutabarat, Syarani and
rice bran1 10.0 10-20 1.30 n.s n.s Smith (1986)
Grass carp Static water 112 Rice bran: WH leaf meal, groundnut 50 6.53 2.58 108.9 2.79 Murthy and Devaraj (1990)
outdoor cement groundnut cake cake, rice bran and fishmeal
cistern (50:50) (50:21:15:10)
Common Static water 112 Rice bran: WH leaf meal, groundnut 50 3.13 3.21 107.7 3.68 Murthy and Devaraj (1990)
Floating aquatic macrophytes – Water hyacinths

carp outdoor cement groundnut cake cake, rice bran and fishmeal
cistern (50:50) (50:21:15:10)
Common Cages in lake 70 No control diet WH meal, fishmeal, corn meal and 2.5 10-20 1.61 n.s n.s Hutabarat, Syarani and
carp rice bran1 10.0 10-20 1.51 n.s n.s Smith (1986)
Rohu Indoor static glass 42 Fishmeal based diet 20 and 40% of total dietary 27 0.20 2.50 79.9 2.31 Hasan, Moniruzzaman and
aquaria (40% protein) protein from fishmeal replaced by 54 0.21 2.21 70.6 2.36 Omar Farooque (1990)
WH leaf meal
Rohu Indoor static glass 77 Fishmeal based diet 25 and 50% of total dietary 30 3.50 0.90 63.7 2.98 Hasan and Roy (1994)
aquaria (30% protein) protein from fishmeal replaced by 60 3.50 0.79 65.3 3.31
WH leaf meal
Stinging Indoor static glass 14 Minced meat (87% Minced meat and WH leaf meal 50 10.5 -3 n.s 2.2 Niamat and Jafri (1984)
catfish aquaria of diet) (37:50)
Java barb Cages in lake 70 No control diet WH meal, fishmeal, corn meal and 2.5 10-20 0.96 n.s n.s Hutabarat, Syarani and
rice bran1 10.0 10-20 1.44 n.s n.s Smith (1986)
Sepat rawa Cages in lake 70 No control diet WH meal, fishmeal, corn meal and rice 2.5 10-20 1.36 n.s n.s Hutabarat, Syarani and
bran1 10.0 10-20 0.93 n.s n.s Smith (1986)
Matrincha Suspended cloth pond 90 Fishmeal, wheat Two levels of WH meal incorporated by 9.5 1.5 1.0 125.0 1.7 Saint-Paul, Werder and
connected to a water meal and corn replacing similar amount of corn meal 18.5 1.5 0.9 112.5 1.8 Teixeira (1981)
treatment plant meal (27:35: 28) from the control diet.

African Plastic tanks 70 Fishmeal based WH meal replaced fishmeal at four 10 1.26 0.64 76.2 3.5 Konyeme, Sogbesan and
catfish diet (35% protein) levels 20 1.26 0.67 79.8 3.3 Ugwumba (2006)
30 1.26 0.63 75.0 3.4
40 1.26 0.54 64.2 3.6
1
One test diet contained 2.5% WH meal with fishmeal, corn meal and rice bran in the ratio of 35.0:10.0:52.5. The other test diet contained 10% WH meal with 7.5% reduction of rice bran
2
SGR not reported. Consumption of control diet was 72%, while it was 70% for test diet. The performance in terms of weight gain was similar for fish fed control and test diet
3
SGR not reported. Fish fed water hyacinth diet registered around 20% gain in live weight in contrast to the group fed control diet where weight gain was about 8%
59
60 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

4.5.3 Composted and fermented forms


Composting or fermentation are techniques often used to reduce water hyacinth into
forms utilizable for feeding livestock.
Composting is one of the most widely used processing techniques to prepare water
hyacinth for use as a fertilizer or fish feed (Figure 4.3). A large quantity of inorganic
nitrogen and phosphorus accumulates in the roots of water hyacinth, which makes it
suitable as a compost or inorganic
Figure 4.3 fertilizer. However, a major
Two farmers carrying dry water hyacinth to the pond side for
problem with the use of water
preparation of compost pit (Mymensingh, Bangladesh)
hyacinth meal in fish diets is its
relatively high crude fibre content.
Fish do not appear to produce
cellulase directly (Buddington,
1980) and their ability to maintain
a symbiotic gut flora capable of
hydrolyzing cellulose is limited.
Fish often poorly accept water
hyacinth leaf meal in pelleted diets.
This has been identified as one
of the major contributory factors
for the reduced growth responses
of major carp (L. rohita) fry by
Hasan, Moniruzzaman and Omar
Farooque (1990). Composting has
been reported to increase the nutritive value and acceptability of water hyacinth.
Edwards. Kamal and Wee (1985) made a comparison of the proximate composition
of composted water hyacinth and dried water hyacinth meals and observed that
while the crude protein levels were similar, the crude fibre and crude fats levels were
approximately halved and the ash content approximately doubled by the composting
process.

Preparation and use of composted water hyacinth


The most commonly used method for compost preparation is the Chinese method of
surface continuous aerobic composting. Edwards. Kamal and Wee (1985) described
the method as follows. Whole water hyacinth plants are cut into 2-3 cm pieces by a
rotary chopper and sun-dried to an ambient equilibrium moisture content of about 20
percent on a platform elevated above the ground to facilitate drying. Compost is made
by mixing dried and freshly chopped water hyacinth to give an initial pile moisture
content of 65-70 percent; the mixture is made into a pile 2.5 m (length) x 2 m (width)
x 1.3 m (height) and perforated bamboo poles are inserted for aeration. The mixture is
turned occasionally to facilitate decomposition. The composting process is completed
within 50 days.
Urea is often added at 2 percent to speed up the decomposing process. In this
process it is suggested that the compost should be prepared by mixing water hyacinth,
cow dung, urea and lime; water hyacinth and cow dung constituting the bulk of the
ingredients while urea and lime are added at 2-5 percent of the total. The ingredients
are kept in an earthen pit and arranged in layers with the top covered by polythene,
paper or banana leaves (Figures. 4.4 and 4.5). Perforated bamboo poles are inserted for
aerobic decomposition. However, compost preparation has been reported to be labour
intensive and farmers are often reluctant to prepare compost for use as fertilizer. A
simple compost preparation technique for use in fish ponds has been developed by
the Mymensingh Aquaculture Extension Project (MAEP, Bangladesh) by using water
hyacinth, cow dung, urea and lime (M.A. Mirza, MAEP pers com. 2004). Freshly procured
Floating aquatic macrophytes – Water hyacinths 61

whole water hyacinths are Figure 4.4


chopped into small pieces A compost pit prepared with water hyacinth and cow dung
and dried for 1-2 days in (Mymensingh, Bangladesh)
sunlight. Sun-dried water
hyacinth containing about
15-20 percent moisture
is mixed with cow dung,
lime and urea in the ratio
of 88:10:1:1 (water hyacinth:
cow dung: lime: urea). The
ingredients are not kept in
layers as traditionally used
but are thoroughly mixed.
The mixture is kept for
decomposition in a pit near
the pond side. The mixture
is re-mixed every seven days
to facilitate decomposition.
The minimum area of the
compost pit is 1 ft2, with a recommended depth of 4 ft to hold 70 kg of compost. The
composting process is normally completed within two months. The recommended rate
of compost application as suggested by MAEP is 18 000 kg/ha/year.
Compost is traditionally used as fertilizer in fish ponds in many Asian countries.
Reports on its use as a fish feed are rather limited, however. Composted water
hyacinth was evaluated as fish feed in pelleted diet for Nile tilapia by Edwards, Kamal
and Wee (1985). These authors prepared four test diets by incorporating 25, 50, 75
and 100 percent of composted water hyacinth meal in a control diet that consisted
of a conventional pelleted tilapia feed (32 percent protein). Good growth and feed
utilization efficiencies were obtained with diets containing up to 75 percent composted
water hyacinth, with no significant reduction in fish performance compared to the
control diet. The specific growth rates varied between 1.96 and 2.15 for test diets while
the SGR for control diet was 1.99. The FCR was between 2.18 and 2.57 for the diets
with compost and 2.63 for the control.
Similarly Hutabarat, Syarani and Smith (1986) reported good growth by using
composted water hyacinth in
a pelleted feed for Java tilapia, Figure 4.5
Java barb and common carp in View of a compost pit in a corner of a pond
cage culture. However, these (Mymensingh, Bangladesh)
authors used only 10 percent as
their maximum inclusion level.
Edwards (1987) reported that
good results were obtained in
China by composting water
hyacinth with silkworm
faeces (or animal manure) and
quicklime, or by composting
the chopped water hyacinth
with a small amount of salt or
saccharified yeast.
The in situ decomposition
of water hyacinth and its
efficacy was studied by
Mishra, Sahu and Pani (1988)
62 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

in rearing ponds for Indian major carp (rohu, catla and mrigal) fingerlings. Fresh water
hyacinth was applied at 300 kg/month/0.2 ha pond (1 500 kg/ha/month). The water
hyacinth was killed in situ by using an aqueous solution of 2,4-D sodium salt and was
allowed to decompose and disintegrate in the pond. The ponds were stocked at a rate
of 3 000 fingerlings per ha (600/pond) and reared for twelve months. The addition of
water hyacinth increased the fish production by about 52 percent as compared to the
control pond where no additional input was provided. A net increase of 64.7 kg of fish
was obtained by using 3 600 kg of fresh water hyacinth. The conversion ratio worked
out to be ~55.7 for fresh hyacinth, while the FCR was about 3.3 on a dry matter basis,
considering that fresh hyacinth contains about 6 percent DM.
Rohu (Labeo rohita) larvae were stocked at 1 million/ha by Sahu, Sahoo and Giri
(2002) under three culture conditions: the application of water hyacinth compost
(8 000 kg/ha), inorganic fertilizer (60 kg/ha), or no manure (control). While the total
nitrogen and phosphate levels of the control treatment were 0.02 and 0.04 g/L, those
in the compost treatment were 0.17 and 0.08 mg/l after 15 days of fertilization. In
the inorganic fertilizer treatment the nitrogen level was elevated to 0.12 mg/l after 15
days but the phosphate level remained at 0.04 mg/l throughout the study period. The
plankton volumes were 1.8, 1.2 and 0.4 ml/45 L in the compost, fertilizer and control
treatments, respectively at the time of stocking of larvae. Significantly (P < 0.01)
higher survival and growth were found in the compost treatment compared to other
treatments.

Preparation and use of fermented water hyacinth


Fermentation is an age-old practice in food processing. In many cases fermentation
has been reported to improve the nutritive value of cereal grains and oilseeds by
increasing their protein efficiency ratio, digestibility and the availability of free amino
and fatty acid contents. During fermentation, nutrient losses may occur as a result of
leaching, destruction by light, heat or oxygen, or microbial utilization (Jones, 1975).
Nevertheless, the loss of nutrients during this process is generally small and there may
even be an increase in the nutrient level through microbial synthesis.
Edwards (1987) reported that water hyacinths were processed in China either
mechanically (soaking, mixing, cutting, or grinding) or biologically for feeding to
grass carp and common carp. The biological processing involved green storage and
fermentation in ditches, tubs, or barrels under anaerobic conditions at 65-75 percent
moisture after cutting into 6 cm strips and sealing by a 15 cm layer of dry grass topped
by a 15 cm layer of moist soil; if the material was too moist it could be sun-dried or
mixed with dry hay before sealing.
A simple fermentative treatment with cow dung and urea was evolved to process
and utilize water hyacinth, as a feed and manure for carp culture by Olah, Ayyappan
and Purushothaman (1990). Water hyacinth leaves were chopped into 5 cm pieces
and mixed with 10 percent cow dung and 2 percent urea. The mixture was then kept
in an airtight polystyrene bag and incubated at room temperature (27-32 ºC). These
authors observed that a period of 2-3 weeks was optimal for cellulose degradation
and to improve the nutritive value of water hyacinth. The crude protein content of
the substrate increased from 13.1 to 18.1 percent of the dry weight during 18 days of
treatment.
Olah, Ayyappan and Purushothaman (1990) fed fermented water hyacinth to catla,
rohu, mrigal, silver carp and common carp in trials conducted in plastic pools for four
(Trial 1) and eight week (Trial 2) periods. The stocking density was 19 and 5/m2 for
Trials 1 and 2, respectively, with daily feeding rates of 50 g/m2. Silver carp and mrigal
showed the best growth rates, followed by rohu. Food conversion ratios of 2.02 and
3.72 were obtained for Trials 1 and 2, respectively. Fermentation of water hyacinth
Floating aquatic macrophytes – Water hyacinths 63

may thus be a simple and efficient treatment for utilizing water hyacinths as a feed or
manure in fish culture without the energy-consuming process of pelletization.
Xianghua (1988) reported on the use of fermented water hyacinth as feed for grass
carp. The plant was harvested, chopped, blended with a small amount of corn flour
and fermented overnight. Good results were obtained in rearing grass carp beyond age
II+.
El Sayed (2003) reported that Nile tilapia (Oreochromis niloticus) fingerlings (1.1 g)
were fed with water hyacinth treated with various processes. Fresh dry hyacinth (FH),
molasses-fermented hyacinth (MF), cow rumen content-fermented hyacinth (RF) and
yeast-fermented hyacinth (YF) were incorporated into nine isonitrogenous (35 percent
CP), isocaloric (450 kcal GE/100 g) test diets, as a substitute for wheat bran at 10 and
20 percent levels. Fish fed the control diet (wheat bran based) exhibited growth, feed
conversion efficiency and production values significantly (P < 0.05) higher than those fed
with water hyacinth based diets. There was no significant difference in the performance
of fish between the fermentation products added at the 10 percent inclusion level. At
the 20 percent inclusion level, the performance of fish was further reduced. Despite this
rather discouraging result, it is interesting to note that significantly lower growth rate
and feed utilization efficiency was found in fish fed with fresh dry water hyacinth than
when fish were fed the fermented water hyacinth treatments.

4.5.4 Other processed forms


There are other processing techniques that are employed to increase the feeding value
of water hyacinth for livestock, such as boiling, mashing and chopping.
For example, Gohl (1981) reported that boiled water hyacinth is used in Southeast
Asia for feeding to pigs. The plants are chopped, sometimes mixed with other vegetable
wastes such as banana stems, and boiled slowly for a few hours until the ingredients
turn into a paste, to which oil cake, rice bran and sometimes maize and salt are added.
The cooked mixture is good for only three days, after which it turns sour. A common
formula is 40 kg of water hyacinth, 15 kg of rice bran, 2.5 kg of fishmeal and 5 kg of
coconut meal.
In China, mashed water hyacinth is used as feed for Chinese carps (Z. Xiaowei, pers.
com. 2003). Fresh water hyacinth is mashed into a liquid form with a high-speed beater
and applied to ponds for carp fingerlings. The mesophyll cells are considered, rightly or
wrongly, similar to phytoplankton. There is an additional means of using mashed water
hyacinth as fish feed: water hyacinth pastes are mixed with rice bran and are fermented
before applying to the pond.
Kumar et al. (1991) evaluated the nutritive value of mashed water hyacinth leaf
for rohu spawn (1.9 mg). Mashed water hyacinth leaves were fed in the form of leaf
extract. The hyacinth extract was prepared by crushing the leaves with water (1:5) in a
heavy-duty mixer. The solution was sieved through a 1 mm mesh to remove the fibrous
material. One or two percent common salt was added to the solution. The experiment
was conducted for 30 days in 40 L glass tanks. Plankton dominated by rotifers and
cladocerans were used as a control treatment. Hyacinth extract was provided at
100 ml/day to the experimental tank containing 120 spawn. The specific growth rate of
rohu spawn fed with mesophyll cells was 8.59 while that for the control was 9.04.
Edwards (1987) reported the efficiency of three processing techniques applied to
water hyacinth for use as fish feed and fertilizer from unpublished research studies
carried out in the Asian Institute of Technology, Thailand. Water hyacinths were added
to a series of earthen ponds stocked with O. niloticus in three forms: fresh whole plants
that decomposed beneath the water in situ; freshly chopped water hyacinth spread on
the surface; and composted water hyacinth. Extrapolated yields of 5 to 6 tonnes/ha/
year were obtained with all three treatments at the same dry matter loading rate of
200 kg/ha/day (about 3 kg TKN/ha/day).
64 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

Bolenz, Omran and Gierschner (1990) suggested the following treatment to avoid
the problem of oxalate crystals (see section 4.5.1). The plants should be chopped into
small pieces; this helps to eliminate trapped air and negate its ability to absorb water.
Then the solid material should be separated from the soluble components in the juice
by pressing and centrifugation. The solid phase will be washed with acid to remove
the acid-soluble calcium oxalate. The juice may be concentrated, dried and used as a
protein source. However, such elaborate treatments will probably not be cost-effective
in preparing fish feeds.

4.5.5 Food conversion efficiency


Food conversion values of diets containing varying inclusion levels of dried water
hyacinth meal tested for different fish species were included in Table 4.3. It can
be observed that the FCR of these test diets varied between >2.0 and <4.0, with
the exception of Nile tilapia (FCR 4.3), as reported by Klinavee, Tansakul and
Promkuntong (1990) and Brycon sp. (FCR 1.7-1.8), as reported by Saint-Paul, Werder
and Teixeira (1981). However, it is difficult to standardize an FCR from the available
data because of the difference in fish species, water hyacinth inclusion levels, rearing
systems and length of rearing. A summary of food conversion ratios for various fish
species fed test diets containing fresh and processed water hyacinth is presented in
Table 4.4. Apart from the pelleted diets containing dried water hyacinth meal, not
much information on FCR for other forms of water hyacinth is available. However,
what is available indicates that an FCR value of 3.0 is a reasonably acceptable level for
fresh or processed water hyacinths.

Table 4.4
Food conversion ratio of fresh and processed water hyacinth for selected fish species
Form of water Incorporation Fish species Fish size (g) FCR Reference
hyacinth level (%) (DM basis)
Fresh 100 Grass carp 38-104 1.54 Riechert and Trede
(1977)
Dried meal as Various (see Various (see Various (see 1.7-4.3 See Table 4.3
pellet Table 4.3) Table 4.3) Table 4.3)
Composted 25-75 Nile tilapia 14.2-17.9 2.18-2.57 Edwards, Kamal and
Wee (1985)
Decomposed 100 Indian major 3.34 Mishra, Sahu and Pani
carps (1988)
Fermented 100 Silver carp 2.02-3.72 Olah, Ayyappan and
and mrigal Purushothaman (1990)
Fermented with 20 Nile tilapia 1.1 1.6 El-Sayed (2003)
molasses

4.5.6 Digestibility coefficients


Several authors have reported the apparent digestibility coefficients of water hyacinth
when fed to carps and tilapia. These varied between species (Table 4.5) and were
influenced by the level of water hyacinth incorporation (Table 4.6). Lin and Chen
(1983, cited by Wee, 1991) noted that protein from water hyacinth was poorly digested
(58.9 percent) by grass carp. Similarly, Riechert and Trede (1977) concluded from their
feeding trial with fresh water hyacinths that only 50-60 percent of the feed consumed
were actually digested by the grass carp. Apparent protein digestibility (APD) of
water hyacinths by Nile tilapia was reported by Pongri (1986, cited by Wee, 1991).
He reported APD values of 49-65 percent and 46-65 percent for dried and composted
water hyacinth when 37.5 percent of water hyacinth was incorporated in the diet. APD
values of water hyacinth leaf meal for Indian major carps (rohu and catla fingerlings)
were reported by Hasan and Roy (1994) and Nandeesha et al. (1991), respectively
(Table 4.6). Digestibility coefficients decreased with increased dietary incorporation of
water hyacinth. For rohu, APD values were 65 and 78 percent for 60 and 30 percent
Floating aquatic macrophytes – Water hyacinths 65

incorporation levels, while for catla it varied between 48 and 74 percent at incorporation
levels from 45-15 percent. In nature, rohu fingerlings feed predominantly on vegetable
debris and microscopic plants while catla are predominantly zooplankton feeders.
Therefore, it is likely that rohu would be able to digest plant materials better than
catla. In an earlier study with rohu fry (mean weight 0.2 g), Hasan, Moniruzzaman
and Omar Farooque (1990) reported the APD of water hyacinth leaf meal to be 55 and
60 percent for 54 and 27 percent levels of dietary water hyacinth inclusion levels. In
contrast to these results, Ray and Das (1994) reported much higher APD value
(94.0 percent) of water hyacinth leaf meal for rohu fry (3.6 g). Similarly high APD
values of water hyacinth for grass carp and common carp fry were reported by Murthy
and Devaraj (1990) (Table 4.5).
From the foregoing discussion, it is difficult to draw any definite borderline between
digestibility coefficients of carps and tilapia. However, it is apparent that digestibility
coefficients are mainly dependent on the level of dietary incorporation. For all practical
purposes, the protein digestibility of water hyacinth may safely be taken as 70-80
percent at 15-30 percent dietary incorporation levels, while it may be around 50-60
percent at incorporation levels of 45 percent or above.
Table 4.5
Summary of apparent nutrient digestibility coefficients of water hyacinth for selected fish species
Form of Fish species Fish size Digestibility coefficient Reference
water (g) (%)
Hyacinth Dry matter Protein Lipid
n.s. Grass carp 58.9 Lin and Chen (1983, cited by
Wee, 1991)
Fresh Grass carp 20-50 50-60 Riechert and Trede (1977)
Dried Nile tilapia 49-65 Pongri (1986, cited by Wee,
1991)
Composted Nile tilapia 46-65 Hertrampf and Piedad-Pascual
(2000)
Leaf meal Rohu 3.5 65-78 Hasan and Roy (1994)
Leaf meal Rohu 0.2 55-60 Hasan, Moniruzzaman and
Omar Farooque (1990)
Leaf meal Rohu 3.6 94 86 Ray and Das (1994)
Leaf meal Catla 23-32 48-74 63-84 Nandeesha et al. (1991)
Leaf meal Grass carp 6.5 89 97 Murthy and Devaraj (1990)
Leaf meal Common carp 3.1 83 98 Murthy and Devaraj (1990)

Table 4.6
Apparent nutrient digestibility coefficients of water hyacinth leaf meal for two carp species at
different dietary incorporation levels
Fish Size Incorporation level of total diet (%) Reference
species (g) 15 30 45 60 15 30 45
Apparent nutrient digestibility (%)
Protein Fat
Rohu 3.5 - 77.6 - 64.5 Hasan and Roy (1994)
Catla 23-32 73.8 59.9 47.9 - 83.9 77.9 63.1 Nandeesha et al. (1991)
67

5. Floating aquatic macrophytes


– Others

Water hyacinth, duckweed and Azolla, the most common and important floating
macrophytes, have been described in sections 2, 3 and 4. This section covers other
miscellaneous floating macrophytes.

5.1 Classification
A list of commonly occurring species is presented in Table 5.1.

Table 5.1
Common and scientific names of some commonly occurring floating aquatic macrophytes
Scientific name Family Common names
Commelina bengalensis Commelinaceae Pakplab; day flower
Enhydra fluctuans Compositae Hinche sak
Enhydra sp. Compositae -
Hydrocharis dubia Hydrocharitaceae -
Ipomoea aquatica (reptans) Convolvulaceae Water spinach
Pistia stratiotes Araceae Water lettuce
Salvinia auriculata Salviniaceae Water fern
Salvinia cucullata Salviniaceae Water fern
Salvinia molesta Salviniaceae Water fern
Salvinia natans Salviniaceae Water fern
Salvinia rotundifolia Salviniaceae Water fern
Trapa bipinosa Trapaceae Water chestnut
Trapa natans Trapaceae Water chestnut

5.2 Characteristics
Like most other macrophytes, these are self-growing plants that are commonly
found in the shallow stagnant waters of tropical and sub-tropical countries. The
most commonly found are water spinach (Ipomoea), water lettuce (Pistia), water fern
(Salvinia spp.) and water chestnut (Trapa spp.).
Water spinach (Ipomoea aquatica) is a floating plant that roots in marshy soil. It is
native to India, SE Asia, and S. China and is commonly eaten as a vegetable (Edwards,
1980).
Water lettuce (Pistia stratiotes) is a free-floating aquatic plant that is found growing
abundantly in tropical and subtropical region of the world. This plant is very common
in lakes, ponds, ditches, irrigation canals, etc. It is reported to be used as a vegetable in
India (Varshney and Singh, 1976).
Water fern (Salvinia spp.) is a perennial free-floating aquatic herb belonging to the
family Salviniaceae. It has five commonly found species (Table 5.1) and has a wide
native range in the neotropics extending from Mexico and the Galapagos Islands
through Central America and most of tropical South America as far as southern Brazil
(Sculthorpe, 1971). In the Indian sub-continent, the plant was introduced through a
route from Brazil to Germany in 1933 (Hadiuzzaman and Khondker, 1993). Reddy and
DeBusk (1985) reported the biomass yield of S. rotundifolia to be 32 tonnes
DM/ha/year in nutrient non-limiting waters in central Florida, USA. Giant salvinia
68 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

(Salvinia molesta) is a free-floating clonal fern and reproduces only vegetatively.


Though the plant can survive a wide range of temperatures (-3 to 43 ºC), optimal growth
occurs at 24 to 28 ºC (Mcintosh, King
Figure 5.1 and Fitzsimmons, 2003). Stands of
Water chestnut plants grown in a floodplain, Rangpur, salvinia double in 2.2 days with
Bangladesh
adequate supply of adequate nutrients.
During periods of high growth, leaf
size decreases and both leaves and
stems fold, doubling and layering
to cover more of the water surface.
This thick plant growth is harmful for
other organisms as it prevents light
penetration, reduces gaseous exchange
and increases biological oxygen
demand.
Water chestnuts (Trapa sp.)
(Figures 5.1 and 5.2) are floating annual
aquatic plants that grow in slow-
moving water up to 5 meters deep and
are native to warm temperate parts of
Asia and Africa.The nut (kernel) of
water chestnuts is eaten by humans
Figure 5.2 in raw or cooked form.
Water chestnut fruits harvested from a floodplain, Rangpur,
Bangladesh
5.3 Production
In South and Southeast Asia, water
spinach is often grown for use as a
vegetable in India, Bangladesh, Hong
Kong (SAR China), Cambodia,
Thailand, Indonesia and Viet Nam,
sometimes in ponds fertilized with
sewage (Figure 5.3). In China,
water lettuce has been reported to
be cultivated with two other aquatic
macrophytes, water hyacinth
and alligator weed (Alternathera
philoxeroides) (Edwards, 1987).
The plants are usually cultivated in
rivulets, small bays, or swamps to
avoid taking up cultivable land and
are usually fed to pigs. Reddy and
DeBusk (1984) reported the biomass yield of water lettuce to be 72 tonnes DM/ha/year
in nutrient non-limiting waters in central Florida, USA. Water chestnuts are grown in
India, China, Indonesia and Bangladesh.

5.4 Chemical composition


The chemical composition of seven floating aquatic macrophytes is presented in Table
5.2. Of these the moisture content varied between 85-94 percent. Water spinach had
the highest crude protein varying between 24-34 percent, higher lipid contents
(2.7-3.9 percent) and low ash (~13 percent) and crude fibre (10.2-12.7 percent) contents.
The other two macrophytes were moderately rich in crude protein (12-20 percent),
high ash (18-26 percent) and crude fibre (12-20 percent) contents. Clearly, water
spinach is nutritionally superior to other macrophytes; its crude protein is comparable
Table 5.2
Chemical analyses of some common floating aquatic macrophytes
Floating macrophytes Moisture (%) Proximate composition1 (% DM) Minerals (% DM) Reference
CP EE Ash CF NFE Ca P
Water fern (Salvinia cucullata), leaf, India 11.6 2.8 18.3 20.4 46.92 n.s. n.s. Ray and Das (1994)
2
Water fern (Salvinia molesta), leaf, India 77.2 16.3 1.1 21.9 18.5 42.23 n.s. n.s. Murthy and Devaraj (1991a)
Water lettuce (Pistia stratiotes), leaf, India 93.62 19.5 1.3 25.7 11.7 41.83 2.352 0.302 Murthy and Devaraj (1991b)
Water lettuce (Pistia stratiotes), leaf, India 11.4 2.7 18.0 20.1 47.8 n.s. n.s. Ray and Das (1994)
Water spinach (Ipomoea aquatica), leaf, Philippines 27.5 3.0 13.2 11.6 44.7 n.s. n.s. Borlongan and Coloso (1994)
Water spinach (Ipomoea aquatica), leaf, Africa 85.0 24.0 2.7 13.3 12.7 47.3 1.20 0.28 Gohl (1981)
Water spinach (Ipomoea aquatica), leaf, Fiji 90.8 34.3 3.9 12.9 10.2 38.7 n.s. n.s. Gohl (1981)
1
CP = crude protein; EE = ether extract; CF = crude fibre; NFE = nitrogen free extract; Ca = calcium; P = phosphorus
2
Floating aquatic macrophytes – Others

Data obtained from Tacon (1987)


3
Adjusted or calculated; not as cited in original publication
69
70 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

Figure 5.3 only to duckweed. Some information


Production of water spinach from Beung Cheung Ek waste
on the amino acid content of various
water lake, Phnom Penh, Cambodia
aquatic macrophytes is contained in
Annex 1.

5.5 Use as aquafeed


Little work has been conducted on
the use of these miscellaneous floating
macrophytes as feed for fish.
A summary of selected experimental
studies on the use of dried leaf meal of
these macrophytes in pelleted diets
for various fish species is presented
in Table 5.3. Dried water spinach leaf
meal has been evaluated as a dietary
These aquatic macropyte are used for both human and/or livestock consumption ingredient/protein source for milkfish
depending on the season and quality of the plants
and water lettuce (Figure 5.4) and
Courtesy of William Leschen
water fern for grass carp, common
carp and rohu. The performance of
test diets containing various inclusion levels of these macrophyte meals were compared
with control diets. The latter comprised fishmeal-based pellets, the traditionally used
rice bran-oil cake mixture, or a mixture of fishmeal, oil cake and cereal by-products.
Apart from rohu, these floating aquaphyte leaf meals (Table 5.3) produced reasonably
good growth (SGR 2.35-3.29 percent) and food conversion (FCR 1.50-2.11). The
performances of milkfish, grass carp and common carp fed various test diets were
slightly better or comparable to those obtained for the control diets. However, it
should be pointed out that Murthy and Devaraj (1991a, 1991b) conducted their feeding
trials with grass carp and common
Figure 5.4 carp in a static experimental setup
A feeding square filled with mixture of grass and chopped consisting of a 20 m2 cement cistern. A
water lettuce offered to the fish as supplemental feed soil bed of 15 cm was provided to each
(Mymensingh, Bangladesh)
cistern and an initial manuring of cow
dung (15 kg per cistern) was carried
out prior to the start of the experiment.
These authors recorded the plankton
volume fluctuating between 0.007 to
0.041 ml/L in the cisterns, indicating
that test fish obtained a part of their
nutrition from the plankton. The
authors noted that the initial manuring
produced the plankton at first but the
faecal matter of fish and unconsumed
food might have been responsible for
the continued plankton production.
Borlongan and Colso (1994) obtained
an SGR of 3.07 percent for milkfish
fry with water spinach leaf meal in a growth trial conducted in a recirculatory rearing
system. However, in this trial, only 15 percent of the total dietary protein (i.e. 6
percent protein in a 40 percent protein diet) was replaced by water spinach leaf meal.
For rohu, Ray and Das (1994) obtained very low SGRs (0.50-0.57 percent) with test
diets containing various inclusion levels of water fern in a growth trial conducted in an
indoor flow-through rearing system (Table 5.3).when compared with a control diet.
Table 5.3
Performance of different fish species to pelleted feeds containing dried floating aquatic macrophytes
Macrophytes/ Rearing system Rearing Control diet Composition of test diet Inclusion Fish size SGR SGR as FCR References
Fish species days level (g) (percent) percent of
(percent) control
Water spinach/ Brackish water 72 Fishmeal based 15 percent of total dietary protein 23 0.3 3.07 100.3 1.50 Borlongan and
milkfish recirculatory diet (40 percent from fishmeal replaced by water Coloso (1994)
system protein) spinach meal
Water lettuce/ Static water 112 Rice bran: Water lettuce leaf meal, groundnut 50 6.5 2.52 106.3 1.84 Murthy and
grass carp outdoor groundnut cake cake, rice bran and fishmeal Devaraj (1991b)
cement cistern (50:50) (27.9 (50:22:14:10)
percent protein)
Water lettuce/ Static water 112 Rice bran: Water lettuce leaf meal, groundnut 50 3.1 3.48 104.5 1.84 Murthy and
common carp outdoor groundnut cake cake, rice bran and fishmeal Devaraj (1991b)
cement cistern (50:50) (27.9 (50:22:14:10)
Floating aquatic macrophytes – Others

percent protein)
Water lettuce/ Indoor flow- 70 Fishmeal: mustard Water lettuce meal incorporated at 15 10.0 0.57 107.5 2.00 Ray and Das (1994)
rohu through system cake: rice bran four levels in control diet by simple 30 0.54 101.9 2.02
(50:35:15) (35 displacement.
percent protein) 45 0.54 101.9 2.08
60 0.50 94.3 2.66
Water fern1/ Static water 112 Rice bran: Water fern leaf meal, groundnut 50 6.5 2.35 99.2 2.11 Murthy and
Grass carp outdoor groundnut cake cake, rice bran and fishmeal Devaraj (1991a)
cement cistern (50:50) (28.0 (50:24:11:11)
percent Protein)
Water fern1/ Static water 112 Rice bran: Water fern leaf meal, groundnut 50 3.1 3.29 110.4 2.11 Murthy and
Common carp outdoor groundnut cake cake, rice bran and fishmeal Devaraj (1991a)
cement cistern (50:50) (28.0 (50:24:11:11)
percent Protein)
1
Salvinia molesta
2
Salvinia sp.
71
72 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

Generally, macrophytophagous fish do not consume these floating macrophytes


readily in fresh form. Water lettuce is one of the least favoured floating macrophytes
for herbivores (Singh et al., 1967). Some scattered reports, however, are available on
the consumption of these macrophytes in fresh form. For example, water lettuce is
used for feeding Chinese carps in China (Z. Xiaowei, pers. com. 2003). Fresh water
lettuce is mashed into liquid form with a high-speed beater and is applied to the pond
for carp fingerlings. Alternatively, mashed water lettuce is mixed with rice bran and
fermented overnight and applied to the pond. Ling (1967, cited by Edwards, 1987)
also reported that water lettuce was chopped into small pieces and used to feed grass
carp and common carp in China. The plant was also reported to be processed, either
mechanically (soaking, mixing, cutting, or grinding) or biologically. The latter involved
green storage and fermentation in ditches, tubs or barrels under anaerobic conditions
at 65-75 percent moisture after cutting into 6 cm strips and sealing by a 15 cm layer of
dry grass topped by a 15 cm layer of moist soil; if the material was too moist it could
be sum-dried or mixed with dry hay before sealing. Another floating macrophyte
(Hydrocharis dubia) is considered a good feed for grass carp in China (Z. Xiaowei, pers.
com. 2003) and is collected or cultivated for feeding carp fingerlings. In oxbow lakes
located in southwestern Bangladesh, water spinach has been found to be one of the
preferred macrophytes for grass carp. In the Mymensingh region of Bangladesh, fish
farmers often give fresh chopped water lettuce and banana leaves as feed for grass carp
and Java barb (Barbonymus gonionotus) in their fish ponds. Both of these herbivores
readily consume these macrophytes and good results have been observed.
Preliminary studies by Mcintosh, King and Fitzsimmons (2003) involved feeding
three month old (3.5 g) Nile tilapia (Oreochromis niloticus) with live giant salvinia
(Salvinia molesta) in a recirculating system, either alone or with a commercial tilapia
feed, compared to a control with commercial feed alone. Results indicated that though
salvinia was consumed by the fish, there was a weight loss during the 23 day culture
period in fish fed salvinia alone. However, some benefit seemed to have been obtained
when salvinia was fed in addition to the commercial diet and these authors speculated
that a longer feeding period might have demonstrated significant effect. These authors
also noted that incorporating dried salvinia as an ingredient in a mixed feed might have
produced a more pronounced effect. The primary interest of this research was the use
of tilapia in weed control (‘biocontrol’).

Table 5.4
Food conversion ratios (FCR) of selected floating aquatic macrophytes to fish
Macrophytes Fish species Fish Food conversion ratio References
size Dry weight Fresh
(g) basis weight basis
Water spinach Milkfish 0.3 1.50 Borlongan and Coloso (1994)
Water lettuce Grass carp 280 Scott and Orr (1970)
Water lettuce Grass carp 50 De Silva (1995)
Water lettuce Grass carp 6.5 1.84 Murthy and Devaraj (1991b)
Water lettuce Common carp 3.1 1.84 Murthy and Devaraj (1991b)
Water lettuce Rohu 10.0 2.00-2.66 Ray and Das (1994)
Water fern1 Grass carp 6.5 2.11 Murthy and Devaraj (1991a)
Water fern1 Common carp 3.1 2.11 Murthy and Devaraj (1991a)
1
Salvinia molesta

The food conversion values of diets containing varying inclusion of dried water
spinach, water lettuce and water fern (S. molesta) tested for various fish species are
summarised in Table 5.4. The FCR of water spinach for milkfish was 1.50. However,
the inclusion level of water spinach meal was only 23 percent and it remains to be
investigated how this might change with increased inclusion levels. The FCR of water
lettuce for grass carp and common carp was 1.84 but it varied between 2.00 and 2.66
Floating aquatic macrophytes – Others 73

for rohu, depending on the level of inclusion. The FCR of water fern for grass and
common carp was 2.11. For both grass and common carps, the level of inclusion of
water lettuce and water fern was 50 percent of the diet.

Table 5.5
Digestibility of selected floating and emergent aquatic macrophytes
Macrophytes Fish Fish Digestibility (percent)1 References
species size
DM CP EE NFE CF GE
(g)
Water lettuce Rohu 3.6 - 91.5 85.4 92.9 - 52.2 Ray and Das (1994)
Water fern2 Rohu 3.6 - 91.4 21.4 66.0 - 64.3 Ray and Das (1994)
1
DM = dry matter; CP = crude protein; EE = ether extract; CF = crude fibre; NFE = nitrogen free extract; GE = gross
energy
2
Salvinia cucullata

It is difficult to standardize the FCR values of these floating macrophytes based


on the available data. For example, the inclusion level of water spinach meal was
only 23 percent in a fishmeal based pelleted diet, while it is apparent that the good
FCR values of water lettuce and water fern obtained for grass and common carps were
partially obtained because of the plankton produced in the cement cistern. Nevertheless,
FCRs of 2.00-2.50 may be reasonably accepted values for dried water lettuce and water
fern when these are used at dietary inclusion levels of up to 50 percent.
Digestibility coefficients of water lettuce and water fern are available only for rohu.
Water lettuce was well digested by rohu with digestibility coefficients varied between
85.4 and 92.9 percent for crude protein, crude lipid and NFE (Table 5.5). For water
fern, crude protein was well digested (91.4 percent) by rohu but the lipid was poorly
digested (21.4 percent). The digestibility of NFE was moderately good (66.0 percent).
75

6. Submerged aquatic macrophytes

Submerged aquatic macrophytes are defined as plants that are usually rooted in the
bottom soil with the vegetative parts predominantly submerged. Many different types
of submerged aquatic macrophytes have been identified globally.

6.1 Classification
Most submerged aquatic macrophytes belong to the families Ceratophyllaceae,
Haloragaceae, Hydrocharitaceae, Nymphaeaceae and Potamogetonaceae. A list of
the most commonly occurring ones is presented in Table 6.1. These macrophytes are
found in various types of water bodies, including estuaries, rivers, lakes, ponds, natural
depressions, ditches, swamps and floodplains. Like other macrophytes, they compete
with phytoplankton for nutrients, decreasing the productivity of the water and causing
hindrance to the movement of fish, irrigation and navigation.

6.2 Characteristics
Submerged macrophytes are distributed all over the world except some very deep and
cold water lakes in polar countries. For example, submerged macrophytes are found
in Asia, mid-eastern Europe, eastern Africa, north and Central America and Australia
and thus have a wide range of environmental requirements in their natural habitats.
Submerged aquatic macrophytes are more commonly found in shallow stagnant
waters. Some comments on environmental effects on the composition of submerged
aquatic macrophytes are given in section 6.4.

Table 6.1
Common and scientific names of various submerged aquatic macrophytes used as fish feed
Scientific name Family Common name
Blyxa lancifolia Hydrocharitaceae Blyxa
Cabomba caroliniana Nymphaeaceae Fanwort
Ceratophyllum demersum/ C. submersum Ceratophyllaceae Hornwort/Coontail
Chara sp. Characeae Chara
Elodea canadensis Hydrocharitaceae Canadian pondweed
E. densa Hydrocharitaceae Brazilian pondweed
E. trifoliate Hydrocharitaceae Pondweed
Haterrauthera limosa Hydrocharitaceae Water stargrass
Hydrilla verticillata Hydrocharitaceae Oxygen weed
Myriophyllum aquaticum Haloragaceae Water milfoil
M. exalbescens Haloragaceae Water milfoil
M. spicatum Haloragaceae Eurasian water milfoil
Najas graminea Hydrocharitaceae Water velvet/ Najas
N. guadalupensis Hydrocharitaceae Water velvet/ Najas
N. marina Hydrocharitaceae Water velvet/ Najas
Ottelia alismoids Hydrocharitaceae Ottelia
Potamogeton crispus Potamogetonaceae Curlyleaf pondweed
P. gramineous Potamogetonaceae Pondweed
P. nodosus Potamogetonaceae Longleaf pondweed
P. pectinatus Potamogetonaceae Sago pondweed
Ruppia maritima Potamogetonaceae Ruppia
Utricularia vulgaria Nymphaeaceae Bladderwort
Vallisneria Americana Hydrocharitaceae Eelgrass
V. spiralis Hydrocharitaceae Eelgrass
76 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

6.3 Production
Production or cultivation techniques have not been developed for most of the
submerged macrophytes, probably because this has not been necessary. However,
some are used as human food and are therefore cultivated. The tip of the shoots of
the Eurasian water milfoil (Myriophyllum spicatum) is eaten as a vegetable in Java
(Indonesia) and is cultivated there (Cook et al., 1974). The leaves of Blyxa lancifolia
are eaten as vegetables in India, where it is one of the most popular vegetables and is
eaten raw with certain kinds of fish. Another submerged plant, Ottelia alismoides,
is also used for human consumption. The entire plant, except the roots, is cooked as
a vegetable. Information on the standing crop of submerged macrophytes is scarce,
except that Boyd (1968) reported that the standing crop value of submerged plants
and algae in lakes in Alabama ranged from 1-4 tonnes/ha. Westlake (1966) reported
net production of submerged macrophytes ranging from 4 to 20 tonnes DM organic
matter/ha/year in fertile ponds.

6.4 Chemical composition


Chemical analyses of some of the common submerged macrophytes used as fish feed
are presented in Table 6.2. Submerged macrophytes generally have a high water content,
which is usually a major deterrent to their harvest and utilization (Edwards, 1980). The
water content of the submerged macrophytes listed varied from 84 to 96 percent. The
water content of hornwort (Ceratophyllum demersum) is particularly high (93-96
percent) and it can thus be described as an ‘absolutely succulent’ type of macrophyte.
The crude protein values of these macrophytes varied between 9 and 22 percent
DM, although most contained levels of 13-15 percent. Most of the submerged plants
contained less than 4 percent lipid, although there were some exceptions, particularly
for oxygen weed. The ash content varied widely from 10 to over 56 percent; however,
most values were between 15 and 30 percent. Fibre contents varied from 7 to 37 percent
but values between 7 and 11 percent were more common.
The apparently wide variations in proximate composition are due to both
interspecific and intraspecific differences in macrophytes. For example, Boyd (1968)
reported crude protein and ash contents of 10.9 and 16.0 percent respectively for
curlyleaf pondweed (Potamogeton crispus), whereas Pine, Anderson and Hung (1990)
reported values of 15.2 and 49.2 percent respectively for crude protein and ash for
the same macrophyte. Similarly, considerable intraspecific variations in nutritional
composition in hornwort, long leaf pondweed (P. nodosus), oxygen weed (Hydrilla
verticillata) and water velvet (Najas guadalupensis) were observed by these authors.
These variations were more pronounced in the case of ash and fibre contents than
protein and lipid. Such intraspecific variations in nutritional composition may also be
attributed to variations in geographic locations, seasonality and environment.
Muztar, Slinger and Burton (1978) recorded a large variation in crude protein
content (7.5-14.9 percent) in Eurasian water milfoil (M. spicatum), simply due to
difference of locations and seasons, although the plant samples were collected from the
same lake in Canada. There is evidence that the crude protein content increases as the
nutrient content of the water in which the plant is grown increases. Pine, Anderson
and Hung (1989) recorded marked variations in proximate composition and acid
detergent fibre of three macrophytes species (sago pondweed P. pectinatus, long
leaf pondweed P. nodosus and Eurasian water milfoil) grown in canals with either
static or flowing water. The greatest differences found were in the levels of dry matter
(DM), nitrogen-free extract, ash, and acid detergent fibre. These major variations in
proximate composition were possibly correlated with the morphological forms that
the plants developed as a response to either static or flowing water conditions. Larger
shoots were produced in these three macrophytes when grown in canals with flowing
water as opposed to static water (Pine, Anderson and Hung, 1989). Furthermore, Pine,
Table 6.2
Chemical analyses of some common submerged aquatic macrophytes
Moisture Proximate composition1 (percent DM) Minerals1 (percent Reference
(percent) DM)
CP EE Ash CF NFE Ca P
Brazilian pondweed (Elodea densa), USA 90.2 20.5 3.3 22.1 29.23 n.s. n.s. Boyd (1968)
Canadian pondweed (E. canadensis), USA 91.12 14.1 1.4 56.5 14.72 1.752 0.362 Pine, Anderson and Hung (1990)
Chara (Chara sp.), fresh, USA 91.6 17.5 1.6 35.8 23.83 n.s. n.s. Boyd (1968)
Chara (C. vulgaris) 91.6 8.8 0.8 28.3 14.0 48.15 n.s. n.s. Tacon (1987)
Curlyleaf pondweed (Potamogeton crispus), USA 88.2 10.9 2.9 16.0 37.23 1.682 0.242 Boyd (1968)
Curlyleaf pondweed (P. crispus), USA4 15.2 1.3 49.2 n.s. n.s. Pine, Anderson and Hung (1990)
Submerged aquatic macrophytes

Eurasian water milfoil (Myriophyllum spicatum), USA4 86.1 21.8 1.7 25.0 7.5 44.05 2.822 0.412 Pine, Anderson and Hung (1989)
Fanwort (Cabomba caroliniana), USA 93.0 13.1 5.4 9.6 26.83 n.s. n.s. Boyd (1968)
Hornwort (Ceratophyllum demersum), India 93.2 13.7 3.1 30.5 7.5 45.25 1.302 0.322 Venkatesh and Shetty (1978b)
Hornwort (C. demersum), India 95.8 12.9 2.6 32.4 9.1 43.0 n.s. n.s. Hajra (1987)
Hornwort (C. demersum), Thailand 16.2 1.5 19.7 8.3 54.35 n.s. n.s. Chiayvareesajja et al. (1988)
Hornwort (C. demersum), USA 94.8 21.7 6.0 20.6 n.s. n.s. Boyd (1968)
Long leaf pondweed (P. nodosus), USA 84.2 11.2 3.6 10.9 21.73 n.s. n.s. Boyd (1968)
Long leaf pondweed (P. nodosus), USA4 92.7 14.6 1.5 45.2 6.5 32.2 n.s. n.s. Pine, Anderson and Hung (1989)
Oxygen weed (H. verticillata), India 89.8 14.6 7.3 21.6 11.1 45.45 4.402 0.282 Venkatesh and Shetty (1978b)
Oxygen weed (H. verticillata), leaf, India 14.1 6.5 19.3 6.9 53.25 n.s. n.s. Ray and Das (1994)
Oxygen weed (H. verticillata), leaf, Sri Lanka 92.3 21.5 11.3 n.s. n.s. De Silva and Perera (1983)
Sago pondweed (P. pectinatus), USA4 91.8 14.6 1.8 40.8 7.7 35.15 n.s. n.s. Pine, Anderson and Hung (1989)
Water velvet (Najas guadalupensis) 15.7 11.6 10.6 27.8 34.35 0.982 0.152 Buddington (1979)
Water velvet (N. guadalupensis), USA 92.7 9.7 3.9 22.7 29.53 n.s. n.s. Boyd (1968)
1
CP = crude protein; EE = ether extract; CF = crude fibre; NFE = nitrogen free extract; Ca = calcium; P = phosphorus
2
Data from Tacon (1987)
3
Cellulose
4
Mean of proximate composition values of weeds collected from flowing and static water
5
Adjusted or calculated; not as cited in original publication
77
78 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

Anderson and Hung (1990) observed marked differences in the proximate composition
of three aquatic macrophyte species (curlyleaf pondweed P. crispus, Canadian
pondweed Elodea canadensis and Eurasian water milfoil) grown in canals having static
and flowing water for three seasons (winter, summer and fall). For example, during the
winter, the ash content in Eurasian water milfoil was 34.6 percent in static water but
43.5 percent in flowing water. These macrophytes also exhibited significant differences
in lipid levels when growing in static or in flowing water, namely 0.5 and 2.0 percent
(curlyleaf pondweed), 0.4 and 2.65 (Canadian pondweed), and 0.55 and 1.8 percent
(Eurasian milfoil).
For all practical purposes, the crude protein content of Brazilian pondweed and water
milfoil may be assumed to be around 20—22 percent, whereas for other submerged
macrophytes it may be taken as 13—16 percent (although some exceptions are shown
in Table 6.2). Similarly, the
Figure 6.1 crude lipid content of most of
Grass carp - a voracious macrophyte feeder the submerged macrophytes
is around 4 percent or below,
except for fanwort and
oxygen weed, which are >5
percent and some individual
analyses for hornwort and
water velvet (Table 6.2).
The extent of intraspecific
variation does not permit
A grass carp harvested from a private fish farm in Mymensingh, Bangladesh species-wise generalizations
Courtesy of M.C. Nandeesha for the ash and fibre contents
of submerged macrophytes.

6.5 Use as aquafeed


A review of the literature indicates that an extensive number of research studies have
been carried out on various submerged macrophytes in different parts of the world.
However, most of these studies concern effective control of submerged macrophytes
by herbivorous fish. Reports are also available on the species preference and
consumption rates of submerged aquatic macrophytes by herbivorous fish. Submerged
aquatic macrophytes are generally soft in nature, moderately rich in protein and are
preferred by different herbivorous fish. In spite of these attractive qualities, only a
limited number of research studies have been carried out on their potential utilization
as fish feed in pond aquaculture. The results of these studies are variable and species
dependent. The most commonly used as fish feed are chara (Chara sp.) hornwort,
oxygen weed (Hydrilla), water velvet (Najas), water milfoil (Myriophyllum) and
pondweeds (Elodea). Most studies were on grass carp (Figure 6.1) and tilapia and
the submerged macrophytes were fed either in fresh form or as a dried meal within a
pelleted diet.

6.5.1 Research studies


A summary of results of selected growth studies carried out on the use of fresh
submerged aquatic macrophytes for fish is presented in Table 6.3. Fresh macrophytes
are generally given to macrophytophagous fish, either whole or after being cut into
small pieces.
In experiments with controlled feeding regimes wherein experimental fish were
fed fresh macrophytes as a complete diet in clear water systems (glass aquaria or fibre
glass tanks), growth responses were either very poor or negative growth was displayed
(Table 6.3). For example, Hajra (1987) reported an SGR of 0.23 percent for grass
carp when hornwort was fed ad libitum in a clear water fibreglass rearing system.
Table 6.3
Performance of tilapia, grass carp and shrimp fed fresh submerged aquatic macrophytes
Macrophytes/ fish species Rearing system Rearing Control diet Feeding rate Fish size SGR SGR as FCR References
days (percent BW/day) (g) (percent) percent of
Fresh Dry control Fresh Dry
weight weight weight weight
Hornwort/ Nile tilapia Circular cages 395 Chicken pellet n.s. 5 21.7 0.331 42.3 n.s. 15.2 Tantikitti et al. (1988)
in lake (19.9 percent
protein)
Hornwort/ Nile tilapia Cages in lake 153 Chicken pellet n.s. 5 16.7 0.952 69.9 n.s. 4.35 Chiayvareesajja et al.
(19.9 percent (1988)
protein)
Submerged aquatic macrophytes

Hornwort/ grass carp Cement cistern 182 - 125 8 12.0 0.94 128.4 10.3 Venkatesh and Shetty
(1978a)
Hornwort/ grass carp Static fibreglass 30 - Ad 14.4 0.23 97.6 4.1 Hajra (1987)
tank libitum
Hornwort/ grass carp Static fibreglass 15 - Ad 52.2 0.21 96.4 4.05 Hajra (1987)
tank libitum
Oxygen weed/ grass carp Cement cistern 182 - 100 10 12.0 1.17 94.0 9.4 Venkatesh and Shetty
(1978a)
Oxygen weed/ grass carp Irrigation tank 126 - Ad 500 0.94 132.0 Keshavanath and
libitum Basavaraju (1980)
Oxygen weed/ grass carp Cement cistern 120 - Ad 3.0 4.27 45.6 Devaraj, Manissery and
libitum Keshavappa (1985)
Chara/ Tilapia zillii Glass aquaria 56 - Ad 17.9 0.15 n.s. n.s. Saeed and Ziebell (1986)
libitum
Najas marina/ Tilapia zillii Glass aquaria 56 - Ad 16.7 0.12 n.s. n.s. Saeed and Ziebell (1986)
libitum
Elodea densa/ Tilapia zillii Glass aquaria 56 - Ad 20.5 Negative growth displayed Saeed and Ziebell (1986)
libitum
Myriophyllum exasbescens/ Glass aquaria 56 - Ad 19.6 Negative growth displayed Saeed and Ziebell (1986)
Tilapia zillii libitum

N. graminea/ shrimp Glass tank 30 Commercial Ad PL50 (0.5 5.26 102.0 n.s. n.s. Primavera and Gacutan
(Penaeus monodon) pellet (40 libitum g) (1989)
percent protein)
Ruppia maritima/ shrimp Glass tank 30 Commercial Ad PL50 (0.5 2.54 49.0 n.s. n.s. Primavera and Gacutan
(Penaeus monodon) pellet (40 libitum g) (1989)
percent protein)
1
SGR of control diet was 0.78 percent, while fish fed no supplementary diet had SGR of 0.55 percent;
2
SGR of control diet was 1.36 percent, while fish fed no supplementary diet had SGR of 1.14 percent
79
80 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

Similarly, poor or negative growth responses were recorded when T. zillii were fed
various submerged macrophytes (Chara sp., N. marina, E. dens and M. exalbescens)
in a clear water static glass aquarium. Poor performances of Nile tilapia were also
recorded by Tantikitti et al. (1988) when fed with fresh hornwort in cage culture. These
authors evaluated fresh hornwort as feed for Nile tilapia and compared its growth and
profitability with chicken pellets and without supplementary feed. In a 14 month trial
in Songkhla lake in Thailand, chicken pellets produced the best growth (weight gain
290 g, SGR 0.78 percent/day), while the performances of fish fed fresh hornwort and
those not provided with any supplementary feed were similar (hornwort: weight gain
65.25 g, SGR 0.33 percent/day; no supplementary feed: weight gain 87.7 g, SGR 0.55
percent/day). Fish fed fresh hornwort did not have any advantage over fish cultured
without any aquatic weed, either in growth or profitability.
Fish reared in clear water static systems tend to consume much less macrophytes
than those reared in cement cisterns and ponds/tanks. Hajra (1987) reported a
hornwort consumption rate of 25 percent BW/day for grass carp in glass aquaria while
the feeding rate used by Venkatesh and Shetty (1978a) for grass carp for the same
macrophyte in cement cisterns was 100 percent BW/day. The variability in growth
responses between clear water indoor static systems and outdoor rearing systems/
ponds might be attributed to the differences in their consumption rates. Moreover,
submerged aquatic macrophytes usually contain about 13-16 percent protein
(Table 6.2). The dietary protein requirement of tilapia and grass carp is much higher
(32-40 percent), which the macrophytes could not generally provide. Therefore, fish
cultured only on a macrophyte diet either lose weight or grow very slowly. The better
growth responses in cement cisterns, earthen ponds or tanks can also be attributed to
the presence of other food organisms such as plankton, benthos, etc.
It is difficult to compare the performances of different macrophytes because of the
variability of rearing systems, experimental duration and fish species. Nevertheless,
grass carp appeared to have performed better when fed oxygen weed than when fed
hornwort (Figure 6.2). Venkatesh and Shetty (1978a) obtained an SGR of 0.94 percent
BW/day for hornwort, while an SGR of 1.17 was recorded for oxygen weed in the
same experimental study. Devaraj, Manissery and Keshavappa (1985) recorded an SGR
of 4.27 percent for grass carp by feeding oxygen weed ad libitum in an experimental
study conducted for 120 days. CIFA (1981) found hornwort to be a poor inducer of
growth, probably due to its poor digestibility.
Attempts have also been made to use dried submerged macrophytes in pelleted feeds
for fish. Drying reduces the moisture
Figure 6.2 content and increases the stability
Farmers carrying mixture of hornwort and and form of macrophytes. However,
oxygen weed in rickshaw van for feeding their the number of studies is extremely
fish (Jessore, Bangladesh)
limited. A summary of the results
of growth studies carried out on the
use of hornwort meal in dry or semi-
moist pelleted feeds for Nile tilapia is
presented in Table 6.4. Test diets were
prepared by using varying inclusion
levels of hornwort meal ranging from
40-98 percent in combination with rice
bran and/ or fishmeal. In these studies
the performances of fish fed the test
diets were sometimes compared with
control diets that consisted of chicken
pellets or commercial fish pellets
containing 16.8-20.7 percent crude
Submerged aquatic macrophytes 81

protein. In all cases where control diets were used the performances of fish fed the
test diets were significantly lower than the control. In some cases the fish fed the test
diets produced growth responses even lower than those given no supplementary feed.
For example, Chiayvareesajja et al. (1988) fed test diets containing various inclusion
levels of hornwort meal and obtained SGRs varying from 1.01 to 1.21 in cages, while
the SGR of the control diet was 1.36 percent and the fish given no supplementary feed
had a SGR of 1.14. It should also be pointed out that the control diets themselves may
have produced sub-optimal growth, as their protein contents varied between 16.8-
20.7 percent, much lower than the optimum requirements of grass carp found when a
complete diet is tested in a clear water system.

6.5.2 On-farm utilization Figure 6.3


Reports on the on-farm utilization Cultivation of watercress, Nasturtium officinale in a
of submerged macrophytes are bamboo frame for feeding of fish in cages (Son La
rather limited. Bala and Hasan Province, Viet Nam)
(1999) reported the efficient on-
farm utilization of submerged
macrophytes in oxbow lakes located
in southwestern Bangladesh. Oxbow
lakes (local name: baors) are semi-
closed water bodies, cut off from
old river channels in the delta of
the Ganges. There are approximately
600 oxbow lakes in southwestern
Bangladesh, with an estimated
Courtesy of M.G. Kibria
combined water area of 5 000 ha.
Many of these oxbow lakes have
been brought under culture-based fisheries management by screening the inlets and
outlets.
Six carp species, i.e. Indian major carps (rohu, catla, mrigal), Chinese carps (silver
carp and grass carp) and common carp, are regularly stocked and harvested almost
throughout the year. The stocking density and species ratios vary widely between
lakes and depend on the water colour and presence of macrophytes in the lake (Hasan
and Middendorp, 1998; Bala and Hasan, 1999). Fishers generally stock more silver
carp in lakes with green water and more grass carp in lakes with a greater coverage
of floating and submerged macrophytes. The most commonly available aquatic
macrophytes in oxbow lakes are water hyacinth (Enhydra fluctuans), water spinach
(Ipomoea aquatica), duckweed (Lemna minor and L. major), oxygen weed, hornwort,
pondweeds (P. crispus and P. nodosus), eelgrass (Vallisneria spiralis), monocharia
(Monochoria hastata), lotus (Nelumbo nucifera) and water lily (Nymphaea spp.). The
most preferred aquatic macrophytes for grass carp in oxbow lakes are water spinach,
duckweed, oxygen weed, hornwort and pondweeds (Potamogeton). Grass carp also eat
the tender leaves of eelgrass. Average stocking densities and yields of each fish species,
grouped by the predominant water colour (green, brown and clear) of 14 oxbow lakes
managed under the Oxbow Lakes Project II are shown in Table 6.5. Green water lakes
are oxbow lakes with distinct algal blooms, as indicated by low Secchi readings, and
also generally have little or no aquatic vegetation. On the other hand brown water
lakes have comparatively more aquatic vegetation. Clear water lakes mostly have a
comparatively high cover of floating and submerged aquatic vegetation. Green water
lakes produce the highest yield of silver carp while a higher yield of grass carp is
recorded in clear water lakes.
On-farm utilization of aquatic macrophytes in cage culture in oxbow lakes in
southwestern Bangladesh (Figure 6.4) has also been observed by the first author of this
82

Table 6.4
Performance of Nile tilapia (Oreochromis niloticus) fed pelleted feeds containing dried hornwort (C. demersum) meal
Rearing system Rearing Control diet Composition of test diet Inclusion Fish size SGR SGR as FCR References
days level (g) (percent) percent
(percent) of control
Cages in lake 153 Chicken pellet Hornwort meal incorporated at nine levels 47.5-97.7 11.7-18.5 1.01-1.211 74.3- 3.87- Chiayvareesajja et
(19.9 percent in combination with pig feed or fishmeal. 89.0 4.10 al. (1988)
protein)
Indoor static 63 No control diet HW meal: rice bran: fishmeal (4:3:1) 57.1 3.1 2.27 - - Chiayvareesajja and
plywood tank Tansakul (1989)
Earthen pond 153 No control diet HW meal: rice bran: fishmeal (4:3:1) 57.1 4.4 1.73 - 4.05 Chiayvareesajja et
al. (1989)
Cages in lake 90 Commercial HW meal: rice bran: fishmeal (4:3:1) 57.1 88.0 0.52 61.5 n.s. Chiayvareesajja,
pellet (20.7 HW meal: rice bran: fishmeal (4:2:2) 50.0 98.2 0.79 93.5 n.s. Wongwit and
percent protein) Tansakul (1990)
Clear water 77 Chicken pellet HW meal, fishmeal and rice bran 40.0 7.2 1.09 69.0 3.70 Klinavee, Tansakul
fibre glass (16.8 percent (40:14.6:40.2) and Promkuntong
tank protein) (1990)
1
SGR of control diet was 1.36 percent, while fish fed no supplementary diet had SGR of 1.14 percent
Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review
Submerged aquatic macrophytes 83

document. Selected fresh submerged, floating and emergent aquatic macrophytes are
used as feed for fingerling rearing in cages floated in oxbow lakes by farmers, with the
help of local NGOs. Grass carp, common carp, Java barb (Barbonymus gonionotus)
and Nile tilapia fry (1.5-2.0 inch) are stocked and reared for about two months
until they attained about 4-6 inch. Indian major carps (rohu and mrigal) are also
occasionally stocked. The stocking rate varies between 1 400 and 1 600 per 8 m3
(2 m x 2 m x 2 m) cage. Two stocking combinations are normally used: grass carp,
common carp and tilapia; or grass carp and Java barb. Grass carp generally form the
bulk (70-75 percent) of the stock. Chopped or whole fresh macrophytes are put into
the cages in the morning, along with 3 kg of a rice bran—wheat bran—oil cake mixture
(7:1:2). Ad libitum feeding or a fixed quantity of 4-5 kg of fresh macrophytes is provided
to each cage every day. The most
commonly used macrophytes Figure 6.4
are: submerged – pondweeds, Mixtures of selected fresh submerged, floating and emergent
oxygen weed, hornwort and aquatic macrophytes are given as feed for fingerling rearing in
cages floated in oxbow lakes in southwestern Bangladesh
eelgrass; floating – duckweed
(Wolffia arrhiza); and emergent -
Monochoria hastata. Pondweeds,
oxygen weed, hornwort and
duckweed are readily eaten by
grass carp, tilapia and Java barb,
whereas the roots and tender
leaves of Monochoria and the
tender leaves of eelgrass are
generally eaten only by grass
carp. Good results are obtained
with grass carp and tilapia/Java
barb. Jagdish, Rana and Agarwal
(1995) and Aravindakshan et al.
(1999) recommended the use of
aquatic macrophytes such as Hydrilla, Najas, Ceratophyllum and duckweeds as food
for grass carp.

Macrophyte preferences
Soft submerged aquatic macrophytes are readily eaten by certain herbivorous fish.
The most commonly fed are hornwort, oxygen weed, water velvet, water milfoil and
pondweeds. The most efficient herbivorous fish is probably the grass carp (known
in the USA as the white amur). Grass carp feed voraciously on submerged aquatic
macrophytes. Several investigations have been carried out to find the consumption
rates and preferences of submerged aquatic macrophytes by this herbivorous fish.
Although grass carp are not specialized feeders and have been reported to consume
over 170 different species of aquatic macrophytes (Redding and Midlen, 1992), they
Table 6.5
Mean stocking densities and yields of six carp species, grouped by the predominant water colour
(green, brown and clear) of 14 oxbow lakes managed under Oxbow Lakes Project II
Stock/ Yield Water Silver Catla Rohu Common Mrigal Grass Total
colour carp carp carp
Stocking Green 1 785 387 519 322 616 216 3 845
density (no/ha) Brown 997 325 740 634 296 345 3 337
Clear 265 197 598 199 247 423 1 929
Yield (kg/ha) Green 317 76 99 73 77 58 700
Brown 174 58 101 52 36 64 485
Clear 25 34 115 33 9.3 86 307
Source: modified from Bala and Hasan (1999)
84 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

were shown to have a preference for certain macrophytes over others. Cassani (1981)
noted that grass carp prefer submerged, rather than floating macrophytes when they
are supplied in fresh form. According to Prabhavathy and Sreenivasan (1977), grass
carp are known to ignore all aquatic vegetation in the presence of oxygen weed.
Venkatesh and Shetty (1978a, 1978b) fed two submerged aquatic macrophytes (oxygen
weed and hornwort) to grass carp and observed that oxygen weed was the most readily
consumed, the whole plant being eaten in the process. In the case of hornwort, these
authors recorded that the smaller fish preferred only the leaves, while the bigger fish fed
readily on the entire plant. In another study, Bhukaswan, Pholprasith and Chatmalai
(1981) reported that grass carp preferred submerged macrophytes such as water velvet
and oxygen weed and floating macrophytes such as water fern. Mitzner (1978) found
that grass carp of approximately 380 g have a preference for Najas and Potamogeton.
The feeding preferences of the blue tilapia Tilapia aurea (weight ranging from
94-176 g) for five aquatic plants were tested by Schwartz and Maughan (1984). These
authors found that the order of preference was (1) Najas guadalupensis and Chara sp.;
(2) filamentous algae (predominantly Cladophora sp.); (3) Potamogeton pectinatus L.;
and (4) P. nodosus.
However, the results of many studies on the preferences of grass carp and their
feeding rates are not in agreement. For example, E. densa, a non-preferred macrophyte
was eaten at the lowest rates in trials in the Pacific Northwest of the USA but proved
to be the first choice and eaten rapidly in trials in Florida (Van Dyke. Lestie and Nall,
1984) thus contradicting other findings that this plant was only moderately preferred
and consumed. Hornwort was quickly eaten in Arkansas and Colorado lakes, but not
in Florida. Similarly, Bonar et al. (1990) recorded that grass carp fed on E. canadensis
from three lakes at significantly different rates, but ate E. densa from two of the sites at
similar rates. The latter authors further observed that the feeding rate of the grass carp
was positively correlated with the concentration of calcium and lignin, but negatively
correlated to the content of iron, silica and cellulose, the most important predictors for
consumption rate being calcium and cellulose.
Hickling (1966), Prowse (1971) and Wiley, Pescitelli and Wike, (1986) hypothesized
that feeding rate and preference in grass carp were primarily influenced by the time
it took the fish to process or ‘handle’ the plant. Its fibre content or the encrustation
on its surface can affect the handling time. The coarseness of macrophytes, due to the
encrustation by calcium carbonate on their external surfaces, makes them unpalatable
(Boyd, 1968). Because grass carp do not digest cellulose, plant cell walls must be
masticated before contents can be assimilated (Hickling, 1966). Wiley, Pescitelli and
Wike (1986) thought that this would increase the handling time of plants high in
cellulose and should lower the preference ranking and the rate of consumption.
Pine, Anderson and Hung (1989) reported the results of a study where triploid
grass carp were presented with three submerged aquatic macrophytes species (sago
pondweed, Eurasian water milfoil, and longleaf pondweed) in outdoor canals with
static and flowing water in winter, spring and summer. During spring and summer, grass
carp showed distinct variation in their preference for aquatic weed types, depending on
their environmental conditions. Plants of all three species produced longer shoots in
canals with flowing water than with static water. The differences in shoot length might
have altered the consumption rate and preference of the fish. Flowing conditions also
had varying effects on the nutritional content of the plants, as shown in proximate
analyses. The preference of triploid grass carp, however, had no correlation with the
proximate analysis variables of the macrophytes. This suggests that accessibility and
ease of mastication were more important in determining preference than the nutritional
quality of the plants. In a further study, Pine, Anderson and Hung (1990) observed
significant variations in feeding preferences and feed efficiencies of one year old grass
Submerged aquatic macrophytes 85

carp for three submerged macrophytes (curlyleaf pondweed, Canadian pondweed and
Eurasian water milfoil) depending on the season (winter, summer and fall) and the flow
of canal water (static and flowing). These authors attributed the differences in feeding
preferences partly to the accessibility of plants to the fish (owing to the difference in
plant stature); plants in static canals did not grow as long as those in flowing canals.
Aquatic macrophyte preferences of grass carp have also been found to be affected
by the ambient temperature. Redding and Midlen (1992) reported that grass carp
consumed more of the softer and more succulent submerged aquatic macrophytes,
such as Elodea, Hydrilla, Myriophyllum and Potamogeton, when water temperatures
were below 12-15 ºC.
The discrepancies in the results of the various studies reviewed above suggest
that ranking plant palatability on the basis of species type alone would be an over-
simplification. Environmental factors and fish size may also play important roles in
determining the macrophyte preferences and consumption rates of grass carp.
Other herbivorous fish are known to consume submerged aquatic macrophytes,
such as tilapia (Tilapia zillii and T. rendalli), Java barb (Barbonymus gonionotus) and
giant gourami (Osphronemus gorami). It has also been reported that the silver barb
(Puntius gonionotus) controlled dense vegetation of Ceratophyllum and Najas from
a 284 ha reservoir in East Java, Indonesia within 8 months of stocking (Schuster,
1952 cited by Edwards, 1980). This author also noted that T. zillii and T. rendalli
are voracious feeders of submerged macrophytes. T. zillii feeds on various types
but shows preferences when feeding choices are offered. For example, Buddington
(1979) reported that T. zillii preferred Najas guadalupensis as a food source to
Lemna, Myriophyllum spicatum and Potamogeton pectinatus. Saeed and Ziebell (1986)
conducted an experimental study by feeding different macrophytes (Chara sp., Najas
marina, Elodea densa and Myriophyllum exalbescens) to T. zillii and observed that
the most preferred macrophyte was Chara followed by N. marina. E. densa and M.
exalbescens. These authors noted that the coarseness of these macrophytes appeared to
have some influence on its consumption by the fish. N. marina has characteristically
sharp-toothed leaf margins. Fish avoided the terminal bushy twigs on which the leaves
are crowded while taking stems and lower leaves, probably because the spines are less
numerous. Similarly, T. zillii avoided the bulky stems of E. densa and fed on the leaves
and soft slender stems, which are easy to grasp and separate. Like grass carp, T. zillii
also showed a diet shift with increase in size. T. zillii over 9.0 cm long were able to eat
macrophytes better than their juveniles. O. gorami is another fish that feeds mainly
on plant leaves and was introduced into irrigation wells in India from Java to control
submerged macrophytes (Edwards, 1980).

Consumption levels
Ad libitum feeding of fresh macrophytes is generally used for herbivorous fish,
although fresh weight feeding rates of 100-150 percent of body weight (BW)/day are
occasionally recommended for grass carp. These empirical feeding rates have probably
been derived from field observations of the consumption rates of different macrophytes
by grass carp, as reviewed below. The consumption rates of oxygen weed and hornwort
for grass carp were reported to be 100-150 percent BW/day (Singh et al., 1967; Bhatia,
1970). Opuszynski (1972) reported that the consumption rates for smaller sized grass
carp were as high as 100-200 percent BW/day. Based on their field observations and
calculations, Shireman and Maceina (1981) suggested four empirical consumption rates
of grass carp for oxygen weed. These were: 100 percent BW/day for grass carp up to
3 kg; 75 percent BW/day for 3-4 kg; 50 percent BW/day for 4-6 kg; and 25 percent
BW/day for >6 kg. Venkatesh and Shetty (1978a, 1978b) used fresh weight feeding rates
of 100 percent and 125 percent BW/day for oxygen weed and hornwort respectively, in
86 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

a growth trial with grass carp. These authors observed that the these restricted feeding
rates might not have been adequate and recommended ad libitum feeding for grass
carp Bhukaswan, Pholprasith and Chatmalai (1981) reported that grass carp (<1.0 kg)
consume water velvet at levels as high as 243 percent and oxygen weed as high as
191 percent BW/day. In contrast, Hajra (1987) reported much lower consumption rates
of hornwort by grass carp. The mean daily dry matter intake per 100 g body weight
was 0.837 g and 0.977 g in small (14.4 g) and large (52.2 g) fingerlings, respectively. The
fresh weight consumption approximated 25 percent of body weight.
Saeed and Ziebell (1986) recorded distinct variation in consumption while feeding
four different submerged macrophytes ad libitum to T. zillii. The consumption rates
were 79 percent, 67 percent, 24 percent and 16 percent BW/day for Chara sp., N.
marina, E. densa and M. exalbescens respectively.

Food conversion rates


Food conversion values of diets containing varying inclusion levels of dried hornwort
meal in pelleted diets fed to Nile tilapia were presented in Table 6.4. The FCR values
varied between 3.7 and 4.1. All these studies were carried out for Nile tilapia only
and the information for other species was not available. The FCR values were very
similar even though the studies were carried out in different rearing systems, e.g. cages,
earthen ponds and fibre glass tanks. However, considering the highly variable growth
responses of Nile tilapia fed hornwort meal, it may not be appropriate to use these
FCR values without further verification.
Food conversion ratios for fresh hornwort and oxygen weed fed to grass carp are
given in Table 6.6. On a fresh weight basis, the FCR of hornwort varied between 96
and 128, while for oxygen weed it varied between 46 and 132. The apparent variation in
FCR values is not surprising, considering the fact that the feeding trials were conducted
in different experimental systems and under varying environmental conditions, using
fish of different sizes. Devaraj, Maniserry and Keshavappa (1985) reported a fresh
weight FCR of 46 for oxygen weed using 3.0 g grass carp in a cement cistern, while
Keshavanath and Basavaraju (1980) obtained an FCR value of 132 for oxygen weed in
an irrigation canal with 500 g grass carp. Therefore, it is difficult to generalize an FCR
value from the available data. Nevertheless, for practical use, the assumption of FCRs
of 100-125 for hornwort and 60-100 for oxygen weed on a fresh weight basis may be
acceptable.

Digestibility
Digestibility coefficients of hornwort, oxygen weed, E. canadensis, Najas spp. and
Ruppia maritima fed to fish and shrimp (Penaeus monodon) are presented in Table
Table 6.6
Food conversion ratios of hornwort and oxygen weed fed to grass carp and Nile tilapia
Macrophytes Fish species Fish Food conversion References
size (g) ratio
Dry Fresh
weight weight
basis basis
Hornwort Nile tilapia 21.7 15.2 n.s. Tantikitti et al. (1988)
Hornwort Grass carp 12.0 10.3 128.4 Venkatesh and Shetty (1978a)
Hornwort Grass carp 14.4 4.1 97.6 Hajra (1987)
Hornwort Grass carp 52.2 4.05 96.4 Hajra (1987)
Oxygen weed Grass carp 12.0 9.4 94.0 Venkatesh and Shetty (1978a)
Oxygen weed Grass carp 3.0 n.s. 45.6 Devaraj, Maniserry and
Keshavappa (1985)
Oxygen weed Grass carp 500.0 n.s. 132.0 Keshavanath and Basavaraju
(1980)
Oxygen weed Grass carp n.s. n.s. 62.0 Sutton (1974)
Table 6.7
Digestibility of five submerged aquatic macrophytes
Macrophytes Fish/prawn species Fish size (g) Digestibility1 (%) References
DM CP EE NFE CF GE
Hornwort Grass carp 12.0 n.s. 51.0 69.0 n.s. 39.0 n.s. Venkatesh and Shetty (1978b)
Hornwort Grass carp 14.4 51.0 74.9 73.0 51.2 38.2 58.3 Hajra (1987)
Hornwort Grass carp 52.2 49.0 72.0 69.2 49.0 36.8 55.6 Hajra (1987)
Hornwort Grass carp n.s. 49.4 n.s. n.s. n.s. n.s. n.s. Lin and Chen (1983, cited by Wee, 1991)
Hornwort Tilapia rendalli n.s. 48-59 n.s. n.s. n.s. n.s. n.s. Caulton (1982)
Oxygen weed Grass carp 12.0 n.s. 81.0 82.8 n.s. 43.0 n.s. Venkatesh and Shetty (1978b)
Oxygen weed Grass carp n.s. 67.9 n.s. n.s. n.s. n.s. n.s. Lin and Chen (1983, cited by Wee, 1991)
Submerged aquatic macrophytes

Oxygen weed Rohu 3.6 n.s. 82.6 42.8 50.0 n.s. 57.1 Ray and Das (1994)
Oxygen weed Etroplus suratensis n.s. 41.3 64.3 67.2 n.s. n.s. n.s. De Silva and Perera (1983)
N. guadalupensis Tilapia zillii 60.2 29.3 75.1 75.9 n.s. n.s. 45.4 Buddington (1979)
N. graminea Penaeus monodon 30-40 40-47 n.s. n.s. n.s. n.s. n.s. Catacutan (1993)
Ruppia maritima Penaeus monodon 30-40 70-76 n.s. n.s. n.s. n.s. n.s. Catacutan (1993)
1
DM = dry matter; CP = crude protein; EE = ether extract; CF = crude fibre; NFE = nitrogen free extract; GE = gross energy
87
88 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

6.7. Digestibility coefficients varied between both macrophyte and fish species. Dry
matter digestibility appears to be in the range of 40-70 percent, although a rather low
value (29 percent) is reported for N. guadalupensis when fed to Tilapia zillii. Apparent
protein digestibility (APD) varied between 64-83 percent with the exception of the
51 percent APD for hornwort reported by Venkatesh and Shetty (1978b) for grass
carp.
Crude lipid digestibility coefficients varied between 67 and 83 percent (Table 6.7)
with the exception of the 43 percent lipid digestibility of oxygen weed reported for
rohu. The digestibility of NFE of hornwort for grass carp was 49-51 percent and
that of oxygen weed for rohu was 50 percent. Data on the crude fibre digestibility of
hornwort and oxygen weed was available only for grass carp and varied from 37 to
43 percent.
The wide variability in the digestive efficiency of different macrophytes can partly
be attributed to the variation in experimental procedures and techniques employed in
the studies reviewed. In addition, variation in chemical composition and the physical
characteristics of the plants influences digestibility (Buddington, 1979). Nevertheless,
for practical purposes, the dry matter, protein, lipid and carbohydrate digestibility may
be taken as 40-60 percent, 60-80 percent, 70-80 percent and 50 percent respectively for
these common submerged macrophytes.
89

7. Emergent aquatic macrophytes

Emergent aquatic macrophytes are defined as plants that are rooted in shallow water
with vegetative parts emerging above the water surface. It is thought that emergent
macrophytes are the most particularly productive of all aquatic macrophytes since they
make the best use of all three possible states—with their roots in sediments beneath
water and their photosynthetic parts in the air (Westlake, 1963). Westlake (1966)
reported the net yield of emergent macrophytes to range from 35 to 85 tonnes
DM/ha/year in fertile ponds.

7.1 Classification
There are many different types of emergent macrophytes commonly found in the
shallow stagnant waters of tropical and sub-tropical countries of the world. A list of
commonly occurring species is presented in Table 7.1. Most of these macrophytes grow
naturally; some, however, are used for human consumption and are cultivated.

Table 7.1
Common and scientific names of commonly occurring emergent aquatic macrophytes
Scientific name Family Common name
Alisma plantago Alismataceae Water plantain
Alternanthera philoxeroides Amaranthaceae Alligator weed
Cabomba aquatica Nymphaceae Aquarium plant
Colocasia chamissonis Araceae Swamp taro
Colocasia esculenta Araceae Aroids/ Taro
Cyperus esculentus Cyperaceae Sedge
Eleocharis dulcis (tuberosa) Cyperaceae Sedge/ Chinese water chestnut
Eleocharis ochrostachys Cyperaceae Sedge/ Chinese water chestnut
Eleocharis plantagenera Cyperaceae Sedge/ Chinese water chestnut
Euryale ferox Nymphaceae Water lily
Hydroryza aristata Gramineae Swimming grass
Jussiaea repens Onagraceae Water primrose
Justicia americana Acanthaceae Water willow
Leersia hexandra Gramineae Rice cut-grass
Monochoria hastata Pontederiaceae Monochoria
Nelumbo nucifera (speciosa) Nelumbonaceae Lotus
Nuphar luteum Nymphaceae Yellow water lily
Nymphaea lotus Nymphaceae Water lily1
Nymphaea rubra Nymphaceae Red water lily
Nymphaea stellata Nymphaceae Blue water lily
Panicum repens Gramineae Torpedo grass
Polygonum hydropiper Polygonaceae Smart weed
Sagittaria sagittifolia Alismataceae Arrowhead
Sagittaria trifolia (sinensis) Alismataceae Arrowhead
Scirpus acutus Cyperaceae Hardstem bulrush-
Scirpus debilis Cyperaceae Weakstalk bulrush
Scirpus mucronatus Cyperaceae Ricefield bulrush
Sium sisarum Apiaceae Skirret
Sparganium americanum Sparganiaceae Bur-reed
Typha latifolia Typhaceae Cat tail/ Reed-mace
1
There are many species of water lily (e.g. Nymphaea lotus, N. nouchali, N. stellata, Victoria amazonica, V.
cruziana); the most commonly found species is N. lotus
90 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

7.2 Characteristics
Arrowhead (Sagittaria spp.) has eight or more underground stems, each with a corm
at the end. S. trifolia grows wild or semi-cultivated in swamps throughout tropical and
subtropical Asia (Ruskin and Shipley, 1976). It is also widely cultivated in China and

Figure 7.1
Lotus (Nelumbo nucifera), an emergent macrophyte grown in a floodplain, Rangpur, Bangladesh

Lotus leaves are generally quite strong and are often used as disposable plates and as wrapping materials in the countries of south Asia. Please
note the support system on the back side (right) of the leave to show its strength

Hong Kong (Herklots, 1972). S. trifolia and other species of arrowheads are cultivated
by Chinese people in many other parts of the world (Cook et al., 1974). It is reported
to be a serious and widespread weed in many countries. However, since it grows
quickly and no special care is needed, it could probably be developed into a useful crop.
There are no yield data but it can be harvested after 6-7 months (Ruskin and Shipley,
1976). The corms of the arrowhead are boiled like potatoes and are eaten by Chinese
and Japanese people with meat dishes. Sium sisarum is another emergent macrophyte
that is cultivated for its edible roots (Cook et al., 1974). These authors also note that
taro (Colocasia esculenta) has a starch filled rhizome that is often eaten. Sedge (Cyperus
esculentus) is widely cultivated for its edible tubers.
The Chinese water chestnut (Eleocharis dulcis) is an emergent aquatic plant that grows
throughout the year. It is an erect stout and slender perennial leafless sedge (Pandey and
Srivastava, 1991a). It has corms or tubers,
Figure 7.2 which are produced in large quantities
Water lily plant covering a substantial part of Lake
on underground rhizomes towards the
Awasa, Ethiopia
end of the growing season. The plant is
widespread from Madagascar to India, SE
Asia, Melanesia and Fiji. Occasionally, it
is used as a wild source of food in Java
and the Philippines. It is cultivated in
China for the high starch content of its
tubers. It commonly grows in swamps
and shallow waters. The yield was said
to be greater than 7 tonnes/ha (Ruskin
and Shipley, 1976), while Hodge (1956)
noted that the yield is much higher, about
18-37 tonnes/ha. Pandey and Srivastava
(1991a) reported the promising potential
of this plant for leaf protein concentrate
(LPC) production.
Courtesy of P.C. Prabu
Emergent aquatic macrophytes 91

Monochoria hastata is a robust, fast-growing perennial herb commonly found in


ponds, lakes and reservoirs. The fresh biomass yield is about 38-39 tonnes/ha/year
(Pandey and Srivastava, 1991b). These authors also reported that this plant also has
promising potential for LPC production.
Lotuses and water lilies are common aquatic macrophytes that grow naturally in
large natural depressions and lakes, and even in small ditches (Figures 7.1 and 7.2).
Lotus flowers have religious significance for Hindus and Buddhists. The lotus is also
used for human consumption and is widely cultivated in China and India, mainly for
its flowers. The fruits, seeds, rhizomes and stems of water lilies are eaten in S. Asia and
India as vegetables and salad.

7.3 Production
Although some emergent aquatic macrophytes are consumed by humans and cultivated
for this purpose, there is very little in the literature about the way this is practised.
However, there are a few reports of experimental observations.
Sutton (1990) cultured Sagittaria subulata for 32 weeks in pans filled with sand
amended with fertilizers and held in an outdoor tank with flowing pond water. The
dry weight of the plants in the highest level of fertilizer (osmocote 35 g/container)
was 69 g/m2. This was 89 percent less than plants collected from a field population (646
±184 g/m2) in the Wakulla River, Florida. The water temperature ranged from 16.5 to
46.5 ºC during this culture period.
Sharma (1981) reported the growth of Typha elephantina in a drain basin (200-
300 m wide, 5 km long) of the Agricultural Farm at Jaipur. The area was divided into
three zones – submerged, marsh and dry zones depending on the moisture contents.
The net annual production in these zones was as follows: dry zone – 1 991 g/m2/year;
marsh zone 2 327 g/m2/year; and submerged zone 3 696 g/m2/year.
Camargo and Florentino (2000) studied the seasonal variations in the biomass
production of the aquatic macrophyte Nymphaea rudgeana in an arm of the Itanhaém
River (São Paulo State, Brazil). In November (13.1 g DW/m2) a gradual increase of
biomass was recorded that reached a maximum in February (163.1 g DW/m2). Then,
the biomass decreased, maintaining low levels until a new growth period. The reduction
of biomass was associated to the development of floating aquatic macrophytes
(Pistia stratiotes and Salvinia molesta) and, subsequently, to environmental factors
(higher salinity values) that were unfavourable to their development. The net
primary production of N. rudgeana was estimated from the biomass data; the annual
productivity value was estimated between 3.02 and 3.82 tonnes/ha/year.

7.4 Chemical composition


Analyses of the chemical composition of several emergent macrophytes are presented
in Table 7.2. The moisture content of these plants varied between 70 and 92 percent.
Generally, the emergent macrophytes have a lower moisture content compared to
floating and submerged macrophytes. On a dry matter basis the reported crude
protein levels varied between 5 and 40 percent, although most were between 10 and
14 percent. Wide variation is seen in the lipid contents, which ranged between 1.0 and 11
percent. In general, ash contents were relatively low (7 to 20 percent) when compared
with other macrophytes. With the exception of taro (C. esculenta) and alligator weed
(Alternanthera philoxeroides) the emergent macrophytes listed in Table 7.2 had high
crude fibre levels (20 to 33 percent). Nitrogen free extracts varied between 37 and 53
percent. Amongst these emergent macrophytes, Sparganium americanum, C. esculenta
and M. hastata were particularly rich in protein; being 23.8 percent, 25.0 percent and
39.5 percent (DM), respectively.
92

Table 7.2
Chemical analyses of some common emergent aquatic macrophytes
Moisture Proximate composition1 (percent DM) Minerals1 Reference
(percent) (percent DM)
CP EE Ash CF NFE Ca P
Alligator weed (Alternanthera philoxeroides) 84.1 15.1 2.5 20.1 15.1 47.2 n.s. n.s. Tacon (1987)
Aroids/taro (Colocasia esculenta), Nigeria 91.8 25.0 10.7 12.4 12.1 39.8 1.74 0.58 Gohl (1981)
Arrow head (Sagittaria latifolia), USA 85.0 17.1 6.7 10.3 27.63 0.832 0.352 Boyd (1968)
Bur-reed (Sparganium americanum) 89.1 23.8 8.3 11.0 20.2 36.7 n.s. n.s. Tacon (1987)
Cat tail (Typha latifolia), USA 77.1 10.3 3.9 6.9 33.23 0.642 0.172 Boyd (1968)
Monochoria (Monochoria hastata), India n.s. 39.5 n.s. 7.0 n.s. n.s. n.s. Pandey and Srivasta (1991b)
Rice cut-grass (Leersia hexandra), Tanzania 70.0 10.1 1.8 10.4 25.6 52.1 n.s. n.s. Gohl (1981)
Sedge/ water chestnut (Eleocharis dulcis), India 81.1 13.9 n.s. n.s. n.s. n.s. n.s. Pandey and Srivasta (1991a)
Sedge/ water chestnut (Eleocharis ochrostachys), Thailand n.s. 5.0 1.0 11.9 29.2 52.9 n.s. n.s. Klinavee, Tansakul and Promkuntong (1990)
Smart weed (Polygonum hydropiper), USA 80.8 11.9 2.4 7.8 26.93 n.s. n.s. Boyd (1968)
Torpedo grass (Panicum repens), Malaysia n.s. 14.0 2.1 13.4 32.6 37.9 n.s. n.s. Gohl (1981)
Water willow (Justicia americana) 85.0 17.6 3.5 16.1 24.0 38.8 0.82 0.12 Tacon (1987)
1
CP = crude protein, EE = ether extract, CF = crude fibre, NFE = nitrogen free extract, Ca = calcium, P = phosphorus
2
Data obtained from Tacon (1987)
3
Cellulose
Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review
Emergent aquatic macrophytes 93

7.5 Use as aquafeed


Although some of these emergent macrophytes are reasonably rich nutritionally,
most are least preferred by macrophytophagous fish. With the exception of one field
observation on the utilization of fresh M. hastata by grass carp fry in cages, information
on their use as fish feed are almost non-existent. Along with some submerged and
floating macrophytes, chopped or whole Monochoria are supplied into the cages in
the oxbow lakes in southwestern Bangladesh for raising fingerlings. The author of this
document has observed that the roots and tender leaves of M. hastata are often eaten
by grass carp fingerlings.
Similarly, information on the use of dried or processed emergent macrophytes as
fish feed is lacking. Venugopal (1980, cited by Shetty and Nandeesha, 1988) found
that it was possible to replace fishmeal partly with leaf powder of taro (C. esculenta)
in feeds for Indian major carp (catla and mrigal) and common carp. In China, alligator
weed is used for feeding Chinese carps (Z. Xiaowei, pers. com. 2003). Fresh alligator
weed is mashed into liquid form with a high-speed beater and applied to the pond
for carp fingerlings. Alternatively, mashed alligator weed is mixed with rice bran and
fermented overnight before application to the pond. Two per cent table salt is added to
eliminate saponin, which is toxic to fish. Alligator weed has also been reported to be
cooked and mixed with rice bran before being fed to all the important cultured carps
in China (Edwards, 1987). In another study, Klinavee, Tansakul and Promkuntong
(1990) used dried sedge/Chinese water chestnut (Eleocharis ochrostachys) leaf meal in
a pelleted feed for Nile tilapia. These authors incorporated 40 percent dried Chinese
water chestnut meal in the diet, in combination with fishmeal (22.25 percent) and
rice bran (29.25 percent), which was used to feed the fish in indoor aquaria. Growth
performance and food utilization was significantly reduced for fish fed the test diet
containing sedge meal when compared with those fed chicken pellets (16.8 percent
crude protein).
95

8. Conclusions

This section provides some overall conclusions from each section of this review.

8.1 Algae
From the studies conducted to date it may be concluded that:
• Only about 10-15 percent of dietary protein requirement can be met by algae
in test diets without compromising growth and food utilization. There is a
progressive decrease in fish performance when dietary incorporation of algal meal
rises above 15-20 percent. Total replacement of fishmeal by algal meal generally
shows very poor growth responses. Apart from commonly observed impaired
growth, the use of algae as the sole source of protein in fish feed can also result in
malformation.
• The poor performance of fish fed diets containing higher inclusion levels of algae
may be attributable to high levels of carbohydrate, of which only a small fraction
consists of mono- and di-saccharides. A preponderance of complex and structural
carbohydrates may cause low digestibility.
• The collection, drying and pelletization of algae require considerable time and
effort and algal cultivation is costly. Cost-benefit analysis is needed before any
definite conclusions on the future application of algae as fish feed can be drawn.
The use of algae as fish feed additives may be limited to the commercial production
of high value fish.

8.2 Azolla
The following conclusions can be drawn:
• Laboratory feeding trials on the use of fresh or dried Azolla as a complete diet
for fish show inconclusive results. Adequate consideration should be given to the
preference of each target fish to particular species of Azolla before they are used
as feed.
• Similarly fresh Azolla as a complete diet for high-density cage culture may not be
economically viable. However, Azolla may be useful in low density and low input
cage culture.
• As fish food in Azolla-fish pond culture, Azolla contributes directly to weight gain
of macrophytophagous fish. At the same time, increased production of fish faeces
from Azolla fodder may be directly consumed by bottom dwellers in addition
to being used as an organic (nitrogenous) fertilizer to increase overall pond
productivity. However, it should be understood that although the contribution of
Azolla to aquaculture is interesting, it alone could not ensure high productivity. It
can be a useful supplement to natural feed in low-input aquaculture.
• The high rates of decomposition of Azolla make it a suitable substrate for
enriching the detrital food chain or for microbial processing such as composting,
prior to application in ponds.
• The results of several studies indicate that rice-fish-Azolla integration increased
the yield of both rice and fish compared to rice-fish culture alone. The likely
reasons for the increase in rice yield are improved soil fertility resulting from the
increased production of fish faeces from Azolla fodder; reduced weed growth; and
a decreased incidence of insects and pests. Fish yields increase through the direct
consumption of Azolla.
96 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

The advantages of rice-fish-Azolla integration may be summarized as follows:


o increase in fish and rice yields;
o decrease in need for inorganic fertilizers and pesticides;
o decrease in incidence of pests and weeds; and
o improvement of soil fertility.

However, the adoption of rice-fish-Azolla integration depends on the attitude and


capacity of the farmers, the capacity of support services, including the Azolla inoculum
availability, and the overall economic feasibility of the system.

8.3 Duckweeds
Duckweeds have received much attention because of their potential to remove
mineral contaminants from wastewater. Definitive information has been published
on the production and chemical composition of these plants, and their environmental
requirements have been clearly determined. Information on the cultivation techniques
of many duckweed species is also available. Due to their rapid growth, attractive
nutritional properties and relative ease of production, duckweeds have generated
renewed interest among fish nutritionists on their use as possible alternative sources of
fish feed. It can be concluded that:
• The results of laboratory and field studies and on-farm utilization of these
macrophytes clearly indicate that duckweed can provide a complete feed package
for carp/tilapia polyculture.
• Successful use of duckweed as fish feed will ultimately depend on the appropriate
integration of duckweed production and aquaculture. A preliminary model on
duckweed-based aquaculture has been developed and tested under experimental
and farming conditions in Bangladesh. However, there is clearly room for fine-
tuning this model. Further research towards optimization of the species mix and
quantification of feed application for sustainable yield may be necessary.
• The production costs of duckweed, whether as a by-product of wastewater
treatment or produced through farming, will ultimately dictate the success of
duckweed-based aquaculture. It must be emphasized that sufficient quantities
of wastewater may not be available throughout the year to support duckweed
production. Therefore, farmers should have the option of using both wastewaters
and fertilizers (both chemical and organic) to produce duckweed. The market
value of potential fertilizers will eventually determine the economic feasibility
of duckweed cultivation. In many countries, including Bangladesh, duckweed
cannot be grown all year round because water bodies dry up in the dry season.
The availability of fish feed/fertilizer during the dry season needs also to be
addressed in a duckweed-based aquaculture model.

8.4 Water hyacinths


A large number of experimental studies have been carried out on the use of fresh or
processed water hyacinths as fish feed. In general, water hyacinths have been proven
to be moderately successful as a fish feed, although the results are variable. Most of the
laboratory studies carried out on the use of water hyacinth as fish feed concentrated
on the use of dried meal in pelleted feeds. The results of these studies indicate that
water hyacinth leaf meal cannot be used as a fishmeal replacer without compromising
growth and food utilization. It has also been noted that, like all other plant ingredients
or non-conventional feedstuffs, high dietary inclusion levels (75 percent or above) of
water hyacinth meal in complete diets is not feasible, as the minimum dietary protein
requirement for most fish species is above 30 percent. Dried water hyacinth leaf meal
contains 20-23 percent and whole meal 13-16 percent crude protein (DM). It also must
be emphasized that complete diets are not generally used in semi-intensive aquaculture
Conclusions 97

practices in most of the developing countries of the world. Nevertheless, it is clear from
the results of many laboratory studies that dried water hyacinth meal has been used
successfully as an alternative to rice bran and/or the rice bran-oil cake mixture that is
traditionally used as fish feed in many developing countries.
In many of the studies, diets with high water hyacinth inclusion levels performed
poorly when compared to fishmeal-based control diets. It must, however, be pointed
out that it was only the direct nutritional benefit of water hyacinth that was assessed
in these controlled aquarium studies. In natural pond systems, the indirect nutritional
value resulting from the production of natural food enhanced by the fertilization of
uneaten feed and fish faeces should not be overlooked.
Although few studies report the successful use of fresh water hyacinth as fish feed,
it is apparent that the use of processed water hyacinth holds much better promise.
However, the question to be answered is: what processing method would be the most
viable alternative? If a comparison between water hyacinth processed by different
techniques is to be made, we must first consider if the use of dried water hyacinth
meal in pelleted diets is feasible under semi-intensive aquaculture in tropical countries.
From the experience of the first author of this document, the milling of dried of water
hyacinth is labour intensive and pelletization would be even more complicated. Will
fish feed manufacturers come forward to use water hyacinth meal to make the feeds
cheaper? It is unlikely. Therefore, it is conjectured that the use of dried water hyacinth
meal in pelleted feed is not a viable option for tropical small-scale semi-intensive
aquaculture. Similarly, the labour-intensive process of drying and milling of water
hyacinth may also discourage farmers to use this ingredient in farm-made aquafeed.
On the other hand, water hyacinth processed by composting or fermentation
provides similar or higher nutritional benefit but is much less labour intensive. The
preparation of water hyacinth paste requires a high-speed blender and the provision of
electricity, however, and may therefore be less attractive for smallholder aquaculture.
Thus, under the current state of knowledge, it is concluded that composted or
fermented water hyacinth used singly or in combination with other traditional dietary
ingredients holds promise as a supplemental feed for use in semi-intensive fish culture
systems where natural food, produced by fertilization, provides a substantial part of
nutrition for fish. The level of its inclusion, when used in combination with other
ingredients, will vary and will depend upon its availability, processing costs, the fish
species in question and the availability of other ingredients in the locality.
It can therefore be concluded that:
• The use of fresh water hyacinth as an aquafeed is unlikely to be successful.
• The use of dried water hyacinth, though having nutritional value, is unlikely to
become viable for use in small-scale aquaculture.
• The use of composted or fermented water hyacinth, however, does hold promise
as a dietary ingredient in aquafeeds for small-scale aquaculture.

8.5 Other floating macrophytes


Floating macrophytes such as water spinach, water fern, and water lettuce are
reasonably rich in protein. However, apart from some general observations (a) in
Bangladesh that fresh water spinach is a preferred macrophyte for grass carp and that
fresh water lettuce is sometimes given to grass carp and Java barb and (b) that mashed
and/ or fermented fresh water lettuce is used in China for feeding carp fingerlings, no
detailed investigations have been carried out on any other qualitative aspects of their
use.
A limited number of research studies on the use of these other floating macrophytes
in pelleted diets for grass carp, common carp and rohu have been carried out. The results
generally indicated that higher inclusion rates were not able to produce good growth
if the feeds were the only source of nutrition in controlled experimental conditions.
98 Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review

However, when used in a natural or semi-natural rearing system, where plankton and
benthos formed a component of the consumption of the fish, diets containing up to
50 percent water lettuce and water fern produced reasonably good growth for grass
carp and common carp. These studies also apparently indicate that the FCRs of these
macrophytes are reasonably good, varying between 2.0-2.5, when used they are as a
dried meal at maximum inclusion level of 50 percent in combination with other proven
dietary ingredients/protein sources.
The information currently available is insufficient to draw any definite general
conclusions on the suitability of these floating macrophytes for fish feeding in small-
scale aquaculture.

8.6 Submerged macrophytes


A large number of research and field studies have been carried out on the utilization of
various submerged aquatic macrophytes. These have included monitoring the growth
performance, food conversion and digestibility of fish fed different macrophytes; the
determination of their nutritional composition; and the consumption and preference
of various macrophytes by herbivorous fish. Most of these studies, however, were
carried out with grass carp and to a lesser extent with tilapia. The macrophyte species
evaluated so far have been oxygen weed, hornwort, pondweeds, Chara, water velvet
and water milfoil.
Submerged macrophytes were fed to fish either in fresh form or as dried meal
in pelleted diets. Reports on other processing techniques designed to improve their
nutritional qualities are not available. The number of studies have been conducted to
evaluate dried submerged macrophytes as fish feed are extremely limited; so far only
dried hornwort meal has been fed to Nile tilapia in a pelleted diet. The results of the
above feeding trials were inclusive. Fish reared on fresh submerged macrophytes as the
only diet in a clear water rearing system generally produced lower growth responses
than fish reared in cement cisterns or ponds where plankton and benthos formed a
substantial part of the nutrition of the fish. Submerged aquatic macrophytes usually
contain about 13-16 percent crude protein (DM) and were therefore unable to support
good growth if used as the only source of dietary protein. However, when submerged
macrophytes were fed to fish in natural or semi-natural rearing system, they supported
moderate to good growth with fresh weight food conversion ratios varying from 60-
125 percent. Most of the fresh submerged macrophytes were well digested by fish, dry
matter and protein digestibility being 40-60 percent and 60-80 percent, respectively.
Consumption rates for different macrophytes varied between different fish species.
Usually, lower consumption rates were recorded for fish reared in a clear water indoor
rearing system. It is also emphasized that the palatability of the same macrophyte species
may vary considerably, depending on the environmental conditions under which they
grow. Consequently, consumption rates may differ for the same macrophytes for the
same fish species owing to variations in environmental conditions, as well as fish size.
A wide range of consumption rates of hornwort and oxygen weed were reported for
grass carp ranging from 25-200 percent, depending on the fish size. Therefore, to
avoid discrepancy, ad libitum feeding may preferably be practised for feeding fresh
submerged macrophytes.
It is difficult to generalize the preference of submerged macrophytes by different
macrophytophagous fish species, because research studies or field observations were
conducted under different environmental conditions and in different parts of the world
using fish of different age groups. However, it is apparent that grass carp generally
prefer fresh soft submerged macrophytes, preferring oxygen weed and Najas over
Potamogeton, Ceratophyllum, Elodea and Myriophyllum. Tender leaves of eelgrass
(Vallisneria) are also eaten by grass carp. Tilapia (T. zillii) feed on Najas, Chara,
Potamogeton, Elodea and Myriophyllum, although it has shown to have preference
Conclusions 99

for Najas and Chara over other macrophytes. The submerged macrophyte preference
of Java barb has not been documented; it generally feeds on Hydrilla, Ceratophyllum
and Potamogeton, although it is likely that it will feed other soft fresh macrophytes as
well.

8.7 Emergent macrophytes


The limited number of observations reported so far is inadequate to draw any
conclusions on the use of emergent macrophytes as fish feed; further studies are
needed.
101

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119

Annex 1
Essential amino acid (EAA)
composition of aquatic
macrophytes

The EAA of some aquatic macrophytes is provided in Table 1. Further information on


the EAA composition of Azolla and on duckweed is contained in Table 2 and Table 3
respectively.
120

Table 1
Essential amino acid composition of some aquatic macrophytes

Aquatic macrophytes CP EAA ( percent of protein) References


(percent) Arg Hist Iso Leu Lys Met Phen Thr Tryp Val Tyr
Alligator weed (Alternanthera philoxeroides) 15.1 2.10* 1.10* 1.50* 1.90* 1.50* 0.60* Trace* 1.60* - 1.80* - Tacon (1987)
Arrowhead (Sagittaria spp.) 18.2 1.10* 0.60* 0.90* 1.70* 1.60* 0.20* Trace* 1.00* - 1.40* - Tacon (1987)
Azolla (Azolla filiculoides) n.s. 6.62 2.31 5.38 9.05 6.45 1.88 5.64 4.70 2.01 6.75 4.10 Buckingham et al. (1978)
Azolla (Azolla pinnata) Bangkok strain n.s. 11.14 2.19 3.64 7.10 5.77 1.27 4.61 2.82 0.23 4.62 - Almazan et al. (1986)
Canadian pondweed (Elodea canadensis), Canada 14.1 6.95 1.35 4.26 7.45 5.68 1.63 4.47 3.76 1.70 5.32 3.48 Muztar, Slinger & Burton (1978)
Curlyleaf pondweed (Potamogeton crispus), Canada 12.9 6.36 1.40 4.89 8.14 5.12 2.72 4.81 3.72 0.31 5.74 3.14 Muztar, Slinger & Burton (1978)
Chara sp., Canada 6.1 3.94 0.82 3.28 5.57 3.77 0.82 3.44 3.61 1.48 4.43 2.13 Muztar, Slinger & Burton (1978)
Duck weed (Lemna minor), Canada 20.0 5.30 1.60 4.75 8.50 5.65 1.50 4.40 4.40 1.15 5.80 2.85 Muztar, Slinger & Burton (1978)
Eelgrass (Vallisneria americana), Canada 18.3 4.26 0.99 4.10 7.16 2.19 1.26 4.92 3.33 1.15 4.70 3.17 Muztar, Slinger & Burton (1978)
Eurasian water milfoil (Myriophyllum spicatum), 12.8 7.04 1.87 5.76 9.92 7.37 2.12 7.54 4.75 0.60 7.37 3.90 Muztar, Slinger & Burton (1978)
Canada
Oxygen weed (Hydrilla verticillata) n.s. 4.18 1.43 3.89 7.16 4.12 1.63 4.61 3.78 - 4.69 3.55 Boyd (1969)
Sago pondweed (Potamogeton pectinatus), Canada 19.7 4.32 1.12 3.55 5.99 6.45 1.02 4.57 3.15 0.92 4.42 2.54 Muztar, Slinger & Burton (1978)
Water hyacinth (Eichhornia crassipes) n.s. 4.55 1.62 3.86 6.78 4.68 1.37 4.09 3.78 - 4.49 2.93 Boyd (1968)
Water lettuce (Pistia stratiotes) n.s. 3.63 1.69 3.99 7.06 5.27 1.35 4.45 3.84 - 4.82 3.19 Boyd (1968)
Water spinach (Ipomoea aquatica) n.s. 3.31 2.66 3.42 6.55 4.56 1.53 5.67 3.92 - 5.27 4.14 Peñaflorida (1989)
Water willow (Justicia americana) 17.6 3.00* 1.10* 2.50* 4.30* 2.80* 0.90* 2.80* 2.30* - 2.90* - Tacon (1987)

Arg = Arginine; Hist = Histidine; Iso = Isoleucine; Leu = Leucine; Lys = Lysine; Met = Methionine; Phen = Phenylalanine; Thr = Threonine; Tryp = Tryptophan; Val = Valine; Tyr = Tyrosine)
n.s. = not stated.
*Values expressed as percent DM basis.
Use of algae and aquatic macrophytes as feed in small-scale aquaculture – A review
Annex 1. Essential amino acid (EAA) composition of aquatic macrophytes 121

Table 2
Essential amino acid composition of Azolla species
Azolla species Amino acids1 ( percent DM)
Arg Hist Iso Leu Lys Met + Cys Phen + Tyr Thr Val
A. microphylla2 1.90 0.47 1.07 2.29 1.62 0.43 2.17 1.13 1.07
A. caroliniana2 1.58 0.40 0.85 1.96 1.34 0.46 1.93 1.03 0.86
A. filiculoides2 1.04 0.28 0.57 1.42 1.04 0.47 1.29 0.68 0.79
A. nilotica2 1.56 0.37 0.84 1.71 1.27 0.52 1.51 0.91 0.81
A. pinnata var. imbricata2 1.43 0.33 0.76 1.79 1.15 0.21 1.57 0.86 0.88
A. mexicana2 1.33 0.32 0.75 1.66 1.06 0.51 1.45 0.85 0.75
A. pinnata var. pinnata2 1.32 0.32 0.81 1.71 0.96 0.23 1.45 0.84 0.97
A. pinnata3 1.15 n.s. 0.93 1.65 0.98 0.52 1.69 0.87 1.18
1
Arg = Arginine; Hist = Histidine; Iso = Isoleucine; Leu = Leucine; Lys = Lysine; Met = Methionine; Phen = Phenylalanine; Thr =
Threonine; Val = Valine; Cys = Cysteine; Tyr = Tyrosine); crude protein levels not stated
2
modified from Cagauan and Pullin (1991)
3
Alalade and Iyayi (2006)

Table 3
Mean essential amino acid values (g/100 g protein) of four species
of duckweed1 compared to FAO reference EAA pattern
Amino acids Mean ± SD FAO reference protein
Arginine 4.54 ± 0.64 -
Histidine 1.78 ± 0.42 -
Isoleucine 3.61 ± 0.37 4.2
Leucine 6.68 ± 0.58 4.8
Lysine 4.01 ± 0.43 4.2
Methionine 0.90 ± 0.15 2.2
Phenylalanine 4.16 ± 0.39 2.8
Threonine 3.12 ± 0.40 2.8
Tryptophan2 - 1.4
Valine 4.39 ± 0.64 4.2
Tyrosine 2.82 ± 0.44 -
1
L. gibba, S. polyrrhiza, S. punctata and W. columbiana
2
Destroyed during analysis
Source: modified from Culley et al. (1981)
123

Annex 2
Periphyton

Rich periphyton communities boost fish production. The distribution of periphytic


fauna shows differences with regard to quantum and seasonal succession. Periphyton-
supported aquaculture systems offer the possibility of increasing both primary
production and food availability for fish; especially those low in the food chain. The
culture of milkfish (Chanos chanos), a very popular cultured species in Indonesia,
Philippines and Taiwan Province of China, is mainly based on periphytic “lab lab”
as food, the production of which is enhanced by organic and inorganic fertilization
(Juliano, 1985). The “acadjas” of West Africa (Welcomme, 1972), the brush parks of Sri
Lanka (Senanayake, 1981) and the “Katha” fisheries of Bangladesh and India (Wahab
and Kibria, 1994) are well-known examples of periphyton-based aquaculture systems.
Dempster, Beveridge and Baird (1993) have reported that Nile tilapia graze more
efficiently on periphyton substrates than on micro-particles in the water column. Algal
biomass is also higher in periphyton systems. Bhaumik et al. (2005) have reported that
richness of periphytic structure in closed wetlands results in higher fish production
(1 570 kg/ha/year) compared to open system (384 kg/ha/year). Lagoons provided with
substrates for periphyton, supports eight times higher algal biomass compared to the
surrounding lagoons (Konan-Brou and Guiral, 1994).
A range of substrate-supported aquaculture systems (Table 1) have been developed
to reduce the cost of feeding fish (Azim et al., 2002a, 2002b; Keshavanath et al., 2002;
Garg, 2005). In these systems additional substrates are provided for the growth
of periphyton, which has positive effects on fish production. The association of
microorganisms, algae and planktonic organisms attached as periphyton serve as food
for fish and also act as an in situ water purifier ensuring better living conditions. Wahab
et al. (1999) have reported 1.8 times higher production of carp kalbaush (Labeo
calbasu) in ponds provided with scrap bamboo as substrate than from ponds without
substrate. Similar results were also observed with rohu (Labeo rohita) (Azim et al.,
2001), Mahseer (Tor khurdee) (Keshavanath et al., 2001) and milkfish (Chanos chanos)
(Jana et al., 2006). Fish yield is linearly correlated with substrate area (Azim et al.,
2004). Garg (2005) has reported that grey mullet (Mugil cephalus), milkfish (Chanos
chanos), pearlspot (Etroplus suratensis) and Nile tilapia (O. niloticus) are suitable
species for periphyton-based brackish water culture systems. Survival and growth of
these four fish were higher in substrate-supported periphyton-based culture systems
compared to the systems without substrate. The provision of additional substrates in
fish culture ponds reduce the use of artificial feed, especially those species that thrive
low in the food web.

Table 1
Various substrates used in periphyton-based culture system
Fish species Culture system Substrate used Reference
Tilapia Monoculture Dense masses of branches Welcomme (1972)
Sarotherodon melanotheron Monoculture Bamboo poles Hem and Avit (1994)
Labeo calbasu Monoculture Scrap bamboo Wahab et al. (1999)
Labeo fimbriatus Polyculture Bamboo, jutesticks Azim et al. (2002a)
Labeo rohita Monoculture Sugarcane bagasse Ramesh et al. (1999)
Cyprinus carpio Monoculture Paddy straw (Eichhornea sp.) Ramesh et al. (1999)
Tor khudree Monoculture Bamboo poles Keshavanath et al. (2002)
FAO

ISSN 2070-7010
FISHERIES AND
AQUACULTURE
TECHNICAL
PAPER

534

Measuring the contribution


of small-scale aquaculture
An assessment

This technical paper presents a global review on the use of aquatic macrophytes as feed
for farmed fish, with particular reference to their current and potential use by small-scale
farmers. The review is organized under four major divisions of aquatic macrophytes: algae,
floating macrophytes, submerged macrophytes and emergent macrophytes. Under floating
macrophytes, Azolla, duckweeds and water hyacinths are discussed separately; the
remaining floating macrophytes are grouped together and are reviewed as ‘other floating
macrophytes’. The review covers aspects concerned with the production and/or cultivation
techniques and use of the macrophytes in their fresh and/or processed state as feed for
farmed fish. Efficiency of feeding is evaluated by presenting data on growth, food conversion
and digestibility of target fish species. Results of laboratory and field trials and on-farm
utilization of macrophytes by farmed fish species are presented. The paper provides
information on the different processing methods employed (including composting and
fermentation) and results obtained to date with different species throughout the world
with particular reference to Asia. Finally, it gives information on the proximate and chemical
composition of most commonly occurring macrophytes, their classification and their
geographical distribution and environmental requirements.

ISBN 978-92-5-106420-7 ISSN 2070-7010

9 789251 064207
I1141E/1/10.09/1700

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