Observations and laboratory experiments are summarised that examine the widely discussed issue of what Gammarus pulex feeds on. On the basis of the observations of the authors and data from the literature it can be noted that G. pulex belongs to the omnivorous organisms, but, depending on the conditions of subsistence, in its rations either plant or animal food can predominate

Borovitskaya, M. P.

English

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Freshwater Biological Association 
FBA Translation (New Series) No. 102 
Titles Biology of the amphipod Gammarus pulex L. in water-bodies of the Leningrad 
district, and its piscicultural importance. 
Author(s) BOROVITSKAYA, M.P. (1956) 
Reference: Trudy vses.gidrobiol.Obshch. 7, 99-122, pp 108-112 
Original language: Russian 
Date of publication of original: 1956 
Translator: J.E.M. HORNE 
Date of publication of translation.: 1977 
Number of pages of translation: 5 pages 
FBA Translations are to be considered as "provisional". As a rule 
they have not been prepared by expert translators, nor have they 
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F.B.A. Translation (New Series) No. 102 
Biology of the amphipod Gammarus pulex L. in water-bodies of the Leningrad 
district, and its piscicultural importance. 
BOROVITSKAYA, M.P. (1956) 
Trudy vses.gidrobiol.Obshch. 7. 99-122, pp 108-112 
Translated by: J.E.M. Home. 
Feeding of gammarids. * 
Until now in the literature we find very contradictory data on the 
feeding of amphipods. Some authors reckon that amphipods feed on animal 
food, others consider them as typical plant-eaters, a third group classifies 
them as omnivorous, and of course there are investigators suggesting that 
gammarids feed on dead animals. 
According to the observations of L.P. Sabaneev (1874), the food of 
gammarids in the Transural lakes consists of algae, infusoria, rotifers, 
cyclopids, daphnids, and also animal and plant residues. Lampert (1900) and 
Sexton (1928) reckon that gammarids feed principally on animal food. Schubert 
(1905), Walter (1906), Thienemann (1912), Wundsch (1922), adhere to the 
opposite opinion, reckoning gammarids to be plant-eating organisms. Kinne 
(1954) reckons that G.duebeni prefers animal food, but for normal development 
it needs green plant food. Ya.A.Birshtein (1940) and S.G. Lepneva (1948) 
indicate that the food of amphipods consists of plant and animal residues. 
Amphipods prefer soft food. Their favourite food appears to be beginning-to-
decompose plants. No less willingly they also devour the corpses of aquatic 
animals, contributing to the cleaning of water-bodies. In mass they gather 
on dead fish, in the aquarium they eat meat. 
In more detail it is necessary to dwell on the works of Haempel and 
Wilier, who conducted detailed studies on the feeding of amphipods. Haempel 
(1908) reckons that the feeding of gammarids appears to be the basic problem 
in breeding them artificially. He investigated the gut contents of 100 
amphipods. The composition of the food was nearly always alike. The great 
part in the gut was found to be plant detritus; it was always possible to 
distinguish also green algae. In some cases he succeeded in identifying 
animal remains in the composition of the food. 
The data of laboratory experiments showed that gammarids appear to be 
omnivorous organisms, but in natural conditions they feed principally on 
plant food, as they live in streams poor in animal food. In different 
circumstances they switch to animal food, but nevertheless it is possible 
to designate them merely as incidental consumers of animal food. 
Translator's Note 
Apart from specific names, the author uses three other terms for 
Gammarus spp. somewhat indiscriminately. These are: 
"rachok" (lit. "small crayfish"), translated as "crustacean". 
"bokoplavy", translated as "amphipods". 
"gammarusy", translated as "gammarids". 
Wilier (1922a) also undertook detailed studies of the feeding 
of amphipods. On the basis of his observations he is led to the conclusion 
that gammarids appear to be plant-eating organisms, since the principal 
role in their feeding is played by tissues of higher plants, while those 
plants are consumed in the middle of which the gammarids live (duckweed, 
elodea, milfoil, Callitriche, Ceratophyllum). Selection of food was not 
observed in the gammarids. Basically they eat living plants, but the 
crustaceans do not ignore also decaying plant residues. In the case of a 
lack of higher plants or an insufficient quantity of them, filamentous algae 
or even animal food can serve as food for gammarids. 
Thus we see that in spite of different opinions on the character of 
the feeding of gammarids nevertheless the majority of authors are inclined 
to reckon them as plant-eating organisms. The, presence in the literature 
of contradictory data on the nutrition of gammarids we explain thus, that 
the investigators worked with different species. 
According to our observations, G.pulex eats green vegetables, 
skeletonises fallen leaves, attacks mosquito larvae. The Baikal-region 
G.lacustris feeds on detritus. We never came to observe these crustaceans 
eating vegetables, leaves or animal food (indeed, our observations on the 
feeding of G.lacustris were carried out in the aquarium and on limited material). 
According to the data of M.Yu.Bekman (1954), G.lacustris appears preeminently 
a detritus-feeding animal. 
The majority of authors based their conclusions about the feeding of 
gammarids only on their observations in natural conditions or in the aquarium. 
For a final solution of the problem of the feeding of gammarids, microscopic 
investigations of the content of their guts are necessary, as was done in their 
time by Haempel and Willer. 
On the basis of our own observations on the feeding of the gammarids 
(G.pulex), we reckon them to be omnivorous organisms which in nature feed for 
the great part on plant food. As plant food the amphipods utilise tissues of 
living plants, but do not ignore decaying vegetable residues. 
It must be said that a restricted choice of food is not characteristic 
of the gammarids, they eat up all organic matter found in the water-body. The 
crustaceans gather in masses both on vegetables and kitchen refuse fallen into 
the stream which passes by the village. We specially threw into the stream 
different raw vegetables to verify whether the gammarids would eat them and 
whether they do not show a preference for some kinds of vegetables. We took 
potatoes, carrots, swedes, beet, cabbage, turnips. We did not notice a 
particular choice of vegetables, but the gammarids less willingly consumed 
swedes and cabbage. 
Amphipods eat up young shoots of plants, and skeletonise leaves which have 
fallen into the water. They eat out the tissue of the leaf, but the veins, even 
the very soft, remain untouched (fig. 1). First of all they eat up the leaves of 
maple and lime, to a lesser degree birch and alder, and they gnaw the winged 
seeds of maple. Into Jordan * fall many oak leaves, but we never observed that 
in winter time the gammarids used them in food. 
* a spring in the Leningrad area. 
In the spring time, when out of the leaves in the water-body there 
remain only oak leaves, the gammarids attempt to utilise them as food, but 
nevertheless not once did we succeed in observing that the leaves of oak were 
completely or even half eaten. Apparently, in view of the significant content 
of tannin, oak leaves are little good as food for amphipods. 
V.Ya.Levanidov (1949) notes that according to chemical composition, 
the fallen leaves, in spite of their poverty in minerals and nitrogenous 
matter, can serve as valuable food for many aquatic invertebrates. 
In the summer of 1950 we carried out the following experiment. In a 
flow-through pond, which was fertilized with willow-herb and also branches of 
alder, were placed the amphipods. Here occasionally were thrown potatoes and 
potato peelings. The crustaceans in this pond successfully bred, while in 
the overwhelming majority they kept to the fertilized zone. This experiment 
shows that amphipods can live in ponds; in the fertilized zone they find 
abundant food. 
Next we carried out the following experiment. Amphipods, brought from 
Ropsha, were divided into several groups and put in jars. Part of the 
amphipods received animal food in the form of mosquito larvae, another part 
fallen leaves of willow, maple and lime, a third group of amphipods lived on 
green food (elodea, vallisneria), a fourth group received as food raw potatoes, 
and lastly, a fifth group - mixed food. There was no soil in any of the jars. 
Observations were made during a month. The experiment showed that the food most 
suited to the gammarids appears to be mixed food or fallen leaves. Among the 
amphipods receiving only mosquito larvae, or only elodea, or only raw vegetables, 
a significant mortality was observed, and they ceased to reproduce. The greatest 
mortality of amphipods we noted with feeding on one animal food. In the jars 
with fallen leaves or mixed food appeared pairs, females and eggs. 
These experiments again emphasize the large role of fallen leaves in the 
nutrition of gammarids, and also this, that in aquaria the gammarids prefer 
plant food to animal. At the same time, they never refuse animal food (beef, 
boiled egg-white, meat of molluscs and frogs). We often happened to observe 
gammarids eating larvae of mosquitoes and mayfly. They attack living larvae 
while often enough the size of the larvae exceeds that of the amphipods. 
Gammarids eat not only live animal food, but do not refuse also corpses 
of animals. Not infrequently we saw gammarids devouring dead gammarids. With 
different baits submerged in the stream (corpses of frogs, dead fish, skinned 
mole carcasses), even after a quarter of an hour, individual gammarids swam 
up to the bait, and after several hours gammarids congregated on the bait in 
such quantities that sometimes it was seen only as a dark moving mass of 
crustaceans. 
In the laboratory we often observed the disappearance of females from the 
jars where the pairs had been transplanted. After moulting they not 
infrequently became victims of the males. Repeatedly we observed that the 
adult amphipods destroyed their young. Evidently, in aquaria, with a shortage 
of other food, the crustaceans eat their like. 
Often in aquaria a strange phenomenon was observed - the eggs suddenly 
began to disappear from the females; sometimes their disappearance took place 
gradually, but more often they disappeared fairly quickly. In natural 
circumstances in the females, apparently, the eggs do not disappear, in as 
much as we did not notice, that in just-caught females the brood chamber was 
not completely filled by eggs. An analogous phenomenon was observed by M. 
Yu.Bekman (1954) in G.lacustris. In the opinion of this author, in captivity, 
especially at low temperatures, the females often eat the eggs from their own 
brood pouches. 
On one occasion, together with the captured amphipods were taken eggs 
of trout. The amphipods were placed in a jar along with the eggs, which very 
quickly perished. After several days all the eggs were eaten, only the shells 
of the eggs remained. On the basis of this one can confidently say that 
amphipods destroy dead eggs. Regretfully, we were deprived of the possibility of 
verifying the attitude of gammarids to living eggs of trout. This problem merits 
special investigation. 
We surveyed the gut contents of 50 amphipods (20 amphipods were 
investigated on 18th February 1951 and 30 on 24th August 1951)* Microscopic 
investigations of the gut contents showed that in the feeding of the amphipods 
plant food predominated. Animal residues were found only in 8 crustaceans; 
apparently these were remains of mayfly larvae. In the guts of the gammarids 
were also found protozoan cysts, green algae, and in large quantity diatom 
algae, and also, besides plant detritus, tissues of higher plants. Differences 
in the feeding of the amphipods caught in winter and summer we did not discover. 
Thus, on the basis of our own observations and data from the literature we 
note that G.pulex belongs to the omnivorous organisms, but, depending on the 
conditions of subsistence, in its rations either plant or animal food can 
predominate. 
References [in this part of the paper only] 
Bekman M.Yu. (1954). Biology of Gammarus lactustris Sars in water-bodies of 
the Baikal region. Trudy baikal.limnol.Sta., 14. 
Birshtein Ya.A. (1940). The order Amphipoda. Zhizn' presnykh vod, 1. 
Lampert K. (1900) Zhizn' presnykh vod. Moskva.Devriena. 
Levanidov V.Ya. (1949). The significance of allochthonous materials as food 
resources in water-bodies, especially in the feeding of Asellus 
aquaticus L. Trudy vses.gidrobiol.Obshch. 1. 
Lepneva S.G. (1948). Amphipoda. In: Ocherki iz zhizni presnovodnykh 
zhivotnykh (ed.E.N. Pavlovskii & S.G. Leoneva). 
Sabaneev L.P. (1874). The Transural lakes. Est.-istor.sb. 'Priroda', 
Kn. 1. 
Haempel O. (1908) Uber die Fortpflanzung und kunstliche Zucht des 
gemeinen Flohkrebses (G.pulex and G.fluviatilis). Allg.Fisch.-Ztg., 
33 Jahrgang, N.F., Bd 23. 
Kinne O. (1954a). Zur Biologie und Physiologie von Gammarus duebeni. 
Lillj.Ztschr. f.wiss.Zool.Bd. 157, H. 3 - 4 . 
Schubert (1905) Die Aufzucht der Salmoniden mit Naturfutter. Osterr. 
Fisch.-Ztg., No. 13. 
Sexton E. (1928) On the rearing and breeding of Gammarus in laboratory 
conditions. Journ.Mar.Biol.Assoc. N.S., v.XV. 
Thienemann A. (1912). Der Gergbach des Sauerlandes. Intern.Rev. d. 
gesamten Hydrobiol. u. Hydrograph. Biol. Suppl., IV-V ser. 
Walter E. (1906) Zur Kultur der Flohkrebse. Fisch.-Ztg., Bd. 9. 
Wilier A. (1922a) Nahrungsuntersuchungen bei niederen Wassertieren. III. 
Nahrungsuntersuchungen bei dem Flohkrebs (G.pulex L.). Ztschr.f. 
Fischerei. v. 21 
Wilier A. (1922b) Untersuchungen zur Kenntnis der Ernahrungsphysiologie von 
Gammarus pulex. Schrift. d. physikal.-okonom.Gesellschaft.Jahrg. 63. 
Wundsch H. (1922). Beitrage zur Biologie von Gammarus pulex. Arch. f. 
Hydrobiol. Bd. XIII. H.3. 
Fig. 1. Leaves of maple and lime, skeletonised by amphipods. 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
Notice 
 
 
Please note that these translations were produced to assist the scientific staff of the 
FBA (Freshwater Biological Association) in their research. These translations were 
done by scientific staff with relevant language skills and not by professional 
translators. 
 
 
 


Biology of the amphipod Gammarus pulex L. in water-bodies of the Leningrad district, and its piscicultural importance [Translation from: Trudy Vsesoyuznogo Gidrobiologicheskogo Obshchestva 7 99-122, 1956]

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Biology of the amphipod Gammarus pulex L. in water-bodies of the Leningrad district, and its piscicultural importance [Translation from: Trudy Vsesoyuznogo Gidrobiologicheskogo Obshchestva 7 99-122, 1956]

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