UseAlageAquaticFeed2009 FAO
UseAlageAquaticFeed2009 FAO
UseAlageAquaticFeed2009 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
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
The views expressed in this information product are those of the author(s) and do not
necessarily reflect the views of FAO.
ISBN 978-92-5-106420-7
© FAO 2009
iii
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.
Contents
Introduction 1
1. Algae 3
Classification 3
Characteristics 4
Production 7
Chemical composition 8
Use as aquafeed 8
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
Introduction
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.
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).
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.
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.
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
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
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.
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).
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
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.
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)
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)
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
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.
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
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.
Figure 3.2
Common duckweed, Lemna minor grown
in a pond (Phu Tho Province, Viet Nam)
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.
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).
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.
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
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
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.
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.
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
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)
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
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.
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
4.1 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
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
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.
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.
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
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.
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.
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.
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
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
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
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
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
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.
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.
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.
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.
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
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
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.
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
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
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.
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.
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.
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.
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119
Annex 1
Essential amino acid (EAA)
composition of aquatic
macrophytes
Table 1
Essential amino acid composition of some aquatic macrophytes
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
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
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.
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