Laboratory tests on feeding interactions and food … et al. 1994, Fanta and Meyer 1998, Grotzner...

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POLISH POLAR RESEARCH 20 4 335-346 1999 Edith FANTA Departamento de Biologia Celular Universidade Federal do Parana, 81531-970 Curitiba, PR, BRAZIL e-mail: [email protected] Laboratory tests on feeding interactions and food preferences of some Antarctic fish from Admiralty Bay, King George Island, South Shetland Islands ABSTRACT: The Antarctic nototheniid fish Notothenia neglecta, N. coriiceps, Trematomus newnesi, T. bernacchii, Lepidonotothen nudifrons, Pagothenia borchgrevinki, Gobionotothen gibberifrons and Pleuragramma antarcticum were caught in the Admiralty Bay during the Ant- arctic summer. Their food acceptance was tested in the laboratory under controlled environmen- tal conditions. In 1000 litre tanks fish, krill, and amphipods were offered as food. Some of the of- fered items were detected, recognized, or accepted as food and ingested; some items were re- jected, showing the capacity offish to make choices. All fish fed on krill; seven fish ingested the amphipod Gondogeneia antarctica but Waldeckia obesa was rejected by all species; N. neg- lecta, N. coriiceps, P. borchgrevinki and T. bernacchii preyed on the fish P. antarcticum. N. neglecta is at the top of this food chain, preying on seven of the species that were offered as food and also performing necrophagy and cannibalism, followed by T. bernacchii and P. borchgre- vinki, which accepted five species as food. The more specialized feeders were T. newnesi, which accepted only two of the offered food items. The trophic interactions for all organisms herein studied were established, indicating a difference between the preferential food of some fish when the offer is unlimited, in artificial conditions, and their reported diet when in the natural en- vironment. Key words: Antarctica, King George Island, fish, feeding, interactions. Introduction Antarctic marine environments and especially some coastal regions with depths between 50 and 100 meters, are characterized by relatively constant low temperature, sometimes variable salinity, and ice cover (Fanta et al. 1995, Hubold 1991, Rakusa-Suszczewski 1993) and seasonal changes in the light levels and

Transcript of Laboratory tests on feeding interactions and food … et al. 1994, Fanta and Meyer 1998, Grotzner...

POLISH POLAR RESEARCH 20 4 335-346 1999

Edith FANTA

Departamento de Biologia Celular Universidade Federal do Parana, 81531-970 Curitiba, PR, BRAZIL e-mail: [email protected]

Laboratory tests on feeding interactions and food preferences of some Antarctic fish

from Admiralty Bay, King George Island, South Shetland Islands

ABSTRACT: The Antarctic nototheniid fish Notothenia neglecta, N. coriiceps, Trematomus newnesi, T. bernacchii, Lepidonotothen nudifrons, Pagothenia borchgrevinki, Gobionotothen gibberifrons and Pleuragramma antarcticum were caught in the Admiralty Bay during the Ant­arctic summer. Their food acceptance was tested in the laboratory under controlled environmen­tal conditions. In 1000 litre tanks fish, krill, and amphipods were offered as food. Some of the of­fered items were detected, recognized, or accepted as food and ingested; some items were re­jected, showing the capacity offish to make choices. All fish fed on krill; seven fish ingested the amphipod Gondogeneia antarctica but Waldeckia obesa was rejected by all species; N. neg­lecta, N. coriiceps, P. borchgrevinki and T. bernacchii preyed on the fish P. antarcticum. N. neglecta is at the top of this food chain, preying on seven of the species that were offered as food and also performing necrophagy and cannibalism, followed by T. bernacchii and P. borchgre­vinki, which accepted five species as food. The more specialized feeders were T. newnesi, which accepted only two of the offered food items. The trophic interactions for all organisms herein studied were established, indicating a difference between the preferential food of some fish when the offer is unlimited, in artificial conditions, and their reported diet when in the natural en­vironment.

Key words: Antarctica, King George Island, fish, feeding, interactions.

Introduction

Antarctic marine environments and especially some coastal regions with depths between 50 and 100 meters, are characterized by relatively constant low temperature, sometimes variable salinity, and ice cover (Fanta et al. 1995, Hubold 1991, Rakusa-Suszczewski 1993) and seasonal changes in the light levels and

336 Edith Fanta

photoperiod (Clarke and North 1991; Eastman 1993; Everson 1984; Fanta et al. 1994; Grotzner and Fanta 1998) that may cause seasonal variability in the available food (Clarke 1985).

As such environmental conditions may have prevailed over the past 20 million years, this lead to effects on the composition of the fish fauna (DeWitt 1970). The Nototheniidae have adapted, being able to occupy a large number of very different ecological niches (Hureau 1994), mainly on the continental shelf or the shelf area surrounding the island groups. They also occur in the Admiralty Bay, a fjord and drainage basin in King George Island (Rakusa-Suszczewski 1993).

Most of the fishes are sedentary (Everson 1984), well adjusted to the local en­vironmental conditions, as in Admiralty Bay (Fanta et al. 1989 a, b; 1995). There the food offer varies along the year. Habitat, food and life style often correlates with special morphological or behavioural adaptation (Ekau 1991, Fanta 1994, Fanta et al. 1994, Fanta and Meyer 1998, Grotzner and Fanta 1998, Meyer and Fanta 1998, Rios and Fanta 1998).

When fish share the same environment, competition for food must be mini­mized for successful survival and expansion of the species. This is often provided by diversification of feeding strategies, as well as the flexibility to change from preferable to available food (Fanta et al. 1994, Fanta and Meyer 1998, Meyer and Fanta 1998, Rios and Fanta 1998). The relation between predators and preys is not only size-dependent (Rios and Fanta 1998) but also circumstantial, and often changes in the presence of different species or according to the availability of shel­ters (Fanta et al. 1994, Fanta and Meyer 1998).

In Admiralty Bay some species share specific areas and consequently resting and feeding grounds (Fanta and Meyer 1998, Skóra 1993). The aim of this study was the experimental evaluation of food acceptance and food preferences. The ap­proach was an analysis of the ability of the fish Notothenia neglecta, N. coriiceps, Trematomus newnesi, T. bernacchii, Pagothenia borchgrevinki, Gobionotothen gibberifrons, Lepidonotothen nudifrons and Pleuragramma antarcticum to detect, apprehend, and ingest fish, krill, and amphipods. The results provide an indication of feeding interactions and feeding preferences of species obtained from the same geographical region when the available food is abundant and search for food is not necessary because the experiments are conducted under controlled conditions in tanks.

Material and methods

The following fish were caught in Admiralty Bay, King George Island, South Shetland Islands, during the summer: Notothenia neglecta Nybelin, 1951 (TL 17.5-30.0 cm); N. coriiceps Richardson, 1844 (TL 16.5-19.5 cm); Lepidonotothen (Lindbergichthys) nudifrons (Lonnberg, 1905) (TL 11.3-15.5 cm); Trematomus

Laboratory tests on feeding and food preferences of some Antarctic fish 337

newnesi Boulenger, 1902 (TL 12.7-15.5 cm); T. bernacchii Boulenger, 1902 (TL 25.0-25.5 cm); Pagothenia borchgrevinki (Boulenger, 1902) (TL 10.0-15.0 cm); Gobionotothen (Notothenia) gibberifrons (Lonnberg, 1905) (TL 13.7-20.0 cm) and Pleuragramma antarcticum Boulenger, 1902 (TL 6.5-7.5 cm). All fish were identified according to Fischer and Hureau (1985) and the names adjusted accord­ing to Kock (1992), except for Notothenia neglecta and N. coriiceps which were maintained as two distinct species.

The following species were offered as food: the fish N. neglecta (TL 8.0-15.0 cm), L. nudifrons (TL 8.5-11.7 cm), T. newnesi and T. bernacchii (TL 10.0-15.0 cm), P. borchgrevinki (TL 10.0-15.0 cm), G. gibberifrons (TL 9.0-17.0 cm) and P. antarcticum (TL 6.5-8.0 cm); the krill Euphausia superba Dana, 1852 (TL 3.5-5.5 cm); the amphipods Bovallia gigantea Pfeffer, 1888 (TL 0.4-0.8 cm), Gondogeneia antarctica (Chevreux, 1906) (TL 0.5-1.3 cm) and Waldeckia obesa (Chevreux, 1905) (TL 0.2-0.3 cm). Krill and amphipods were identified respec­tively by V.Gomes (Instituto Oceanografico, Sao Paulo, Brazil) and by C. De Broyer (Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium).

Fish were caught with gillnets from 40 to 80 m depth, by midwater and bottom trawl and traps. Krill was obtained through midwater trawl and the amphipods with traps, trawl, and hand nets.

All species were maintained in separate tanks or aquaria under controlled envi­ronmental conditions (photoperiod of 20 hours light and 4 hours darkness, light in­tensity between 9 and 15 lux, temperature of 0°C, pH 7.2, and salinity 35 ppt).

In 1000 litre tanks the reaction of each species of fish: (i) to each one of the food items separately; and (ii) to all food items together, was observed from the moment in which the food was made available till the end of feeding activity caused by depletion. At least 10 observations were made for each predator in the presence of each one of the food items of situation (i) and of all food items in situa­tion (ii).

Results

The choice of food was specific. When different potential food items were available, each species reacted in a certain way. The offered item could stimulate the fish to identify it as food or not (Table 1). Through typical feeding movements (Fanta et at 1994; Fanta and Meyer 1998), this food was eventually captured.

Selection occurred at different moments: when the offered food was seen, me­chanically perceived, or chemically evaluated at distance; or chemically when it touched the lips; and also after capture, when it came in contact with the inner part of the mouth. At each one of these moments, the selection could result in accep­tance or rejection of food.

338 Edith Fanta

Table 1 Food acceptance by: Notothenia neglecta, N. coriiceps, Trematomus bemacchii, T. newnesi, Pagothenia borchgrevinki, Lepidonotothen nudifrons, Gobionotothen gibberifrons and Ple-uragramma antarcticum. Offered food: (K) krill Euphausia superba; (A) amphipods, (G) Gondogeneia antarctica, (W) Waldeckia obesa, (B) Bovallia gigantea; (F) fish, (Pa) Pleura-gramma antarcticum, (Tn) Trematomus newnesi, (Tb) T. bemacchii, (Gg) Gobionotothen gibberifrons, (Ln) Lepidonotothen nudifrons, (Nc) Notothenia coriiceps, (Nn) N. neglecta, (Pb) Pagothenia borchgrevinki; (ne) necrophagy; (+) fish performed the action in more than 75% of the experimental situations; (-) fish never performed the action; (grey field) situation

is not presented. For each predator, 10 repetitions of each situation were analysed.

Offered food

Food acceptance N. neglecta detection capture ingestion rejection T. bemacchii detection capture ingestion rejection P. borchgrevinki detection capture ingestion rejection N. coriiceps detection

capture ingestion rejection L. nudifrons detection capture ingestion rejection G. gibberifrons detection capture ingestion' rejection T. newnesi detection capture ingestion rejection P. antarcticum detection capture ingestion rejection

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Laboratory tests on feeding and food preferences of some Antarctic fish 339

Some fish like P. antarcticum, T. newnesi, G. gibberifrons and L. nudifrons were preyed by other fish species, mainly N. neglecta and P. borchgrevinki. How­ever, krill and amphipods, once introduced in the containers with fish, were always recognized as potential food by all species.

All fish fed on krill and the presence of E. superba was highly stimulating to them. Whereas all species fed on amphipods, the detection and acceptance of food was species' dependent: G. antarctica was ingested by all fish except L. nudifrons, and B. gigantea by N. neglecta, T. bemacchii and L. nudifrons. On the other hand, W. obesa was not detected as food by T. bemacchii, P. borchgrevinki and G. gibberifrons. All other species were stimulated by its presence, capturing W. obesa but throwing it out immediately.

Among the fish, P. antarcticum was preyed by N. neglecta, N. coriiceps and P. borchgrevinki. If they were swimming very actively they inhibited the feeding of N. coriiceps. Other small specimens like T. newnesi were preyed by N. neglecta and P. borchgrevinki, G. gibberifrons being ingested by both and also by TV. coriiceps.

N. neglecta is at the top of this food chain, preying on seven species but also ex­ceptionally ingesting small individuals of their own species. The next more intense feeders were T. bemacchii and P. borchgrevinki, followed by N. coriiceps. The most specialised feeder was T. newnesi, which accepted only 2 of the offered food items.

The results of the relations between fishes and the food, concerning its detec­tion, apprehension, and ingestion or rejection are summarised in Table 1 and the in­tensity of predation is summarised in Table 2.

Table 2 Classification of preys according to the decreasing intensity of acceptance, with the num­ber of predator species that preyed on each food item. * - notice that N. neglecta is not

preyed upon but may, eventually, ingest small individuals of their own species.

Decreasing levels of

predation

1. 2. 3. 4.

5.

6.

Food item

Euphausia superba Gondogeneia antarctica Pleuragramma antarcticum Bovallia gigantea Gobionotothen gibberifrons Lepidonotothen nudifrons Trematomus newnesi Notothenia neglecta * Notothenia coriiceps Pagothenia borchgrevinki Trematomus bemacchii Waldeckia obesa

Number of fish species

that ingested the food item

S 7 4 3 3 2 2 0 0 0 0 0

Considering only the accepted food items, one may observe what seems to be a sequence in the preference of certain items upon others. This seems to be based on

340 Edith Fanta

die facility of capture, intensity of visual or chemical stimulation, type of prey movements, shape or colour, or some other criteria, which are undetectable to us. N. neglecta is equally stimulated by krill and by small swimming fish, eating am-phipods as a second choice and small species of fish at the bottom as third choice. Necrophagy occurs when there is no living food available. N. coriiceps is more ac­tive in the presence of krill, being sometimes inhibited by a higher density of swim­ming fish. Their second choice seems to be amphipods. T. newnesi eats mostly am-phipods but is highly stimulated by krill. G. gibberifrons usually eats amphipods, mainly when they are close to the bottom, but is highly stimulated by krill. P. borchgrevinki prefers fish, then krill, and its last choice is amphipods. T. ber-nacchii prefers krill, then amphipods and will eventually take some fish. L. nudifrons prefers krill to amphipods.

Discussion

Feeding behaviour of fish can be related to multiple factors such as food offer, food preferences when the offer varies, morpho-functional capacity of food detec­tion and selection, stomach capacity, inter and intraspecific competition for food, among others (Eastman and DeVries 1985, Fanta et al 1994, Fanta and Meyer 1998, Meyer and Fanta 1998, Rios and Fanta 1998).

Studies on the stomach contents of Antarctic fish from different regions have shown a strong relation between the available food and the ingested food (Moreno 1980, Daniels 1982, Eastman 1985, Foster et al. 1987, Barrera-Oro and Casaux 1990, Casaux et al. 1990, Hubold 1991, Kock 1992, Hureau 1994, Takahashi and Iwami 1997, Rios and Fanta 1998).

On the other hand, it was perceived that fish make some choices when feeding, even rejecting some food items (Fanta et al 1994, Dcen et al. 1997, Fanta and Meyer 1998, Meyer and Fanta 1998). Therefore, the food that is found in the stomach of fish captured from the natural environment often reflects the available but not the preferred organisms. Even food which is considered as usual for fish like N. neglecta and G. gibberifrons, such as ophiuroids, polychaetes, isopods, molluscs or algae (Everson 1984, Fischer and Hureau 1985, Kock 1992, Hureau 1994, Been et al. 1997) were never ingested when food items like amphipods, krill, or fish were available in quantity {personal observ.).

At least during the summer, around 23 fish species live in the Admiralty Bay (Skóra 1993). In the Martel Inlet around 10 Nototheniidae are usually found during the summer (Fanta and Meyer 1998), as well as Channichthyidae, Harpagiferidae, and Bathydraconidae (personal observ.).

In some cases, Antarctic fish taxonomy only adheres to the morphological and meristic characters, often based on small samples. A typical case is that of Noto-thenia coriiceps and N. neglecta. Presently it is considered to be one widespread

Laboratory tests on feeding and food preferences of some Antarctic fish 341

species with different populations which occur around all islands of the Southern Ocean and around the Antarctic Continent (Kock 1992). It is believed that the geo­graphical distance or barriers between populations may have caused some genetic and morphological differences and so, for a while, two species, N. neglecta and N. coriiceps were considered as valid (Fischer and Hureau 1985). On the other hand, we have captured both "species" in the Admiralty Bay. They showed some slight morphological differences, which allowed for their classification according to Fischer and Hureau (1985). Some behavioural differences were also detected by Fanta and Meyer (1998), as well as their food choices in the present tests. There­fore, it is not clear if the individuals herein studied belong to one single species with a high variability in all ethological and morphological characters or if they are really two species, or subspecies. The answer might be possible in the future, if such individuals are analysed from the genetic and molecular biological perspec­tive.

Even considering that the physico-chemical conditions in a container are not identical to those in the natural environment, Antarctic fish have been tested in aquaria for the evaluation of the effects of certain environmental conditions on their organisms (Fanta etal. 1989 a, b, 1995; Fanta 1994), and also to evaluate then-feeding behaviour (Fanta et al. 1994, Fanta and Meyer 1998, Meyer and Fanta 1998, Freiberger and Fanta, unpubl.). Aquarium observations have also greatly contributed to the understanding of trophic interactions on benthos (Arntz and Gallardo 1994). A limited area and the availability of sufficient food in tanks, un­der constant environmental conditions, made it possible to test not only the capac­ity of food detection, but also the morpho-physiological possibility and the will­ingness to catch the detected food item, selecting it according to taste, size, consis­tency, or some other unknown or undetectable criteria.

For that reason, a case study with some of the most common species that share an area in the shallow coastal nearshore region of Martel Inlet in the Admi­ralty Bay can help in understanding of some of the local feeding interactions. Evaluations of the stomach content and feeding habits of some fish have been made in the bay (Linkowski et al. 1983, Rios and Fanta 1998), and in the South Shetlands region (Iken et al. 1997, Takahashi and Iwami 1997), but through the tests in tanks herein conducted it is possible to detect if some of the food items are preferred or rejected when enough food is available and therefore the fish are allowed to make choices.

Size selective predation is important (Ellis and Gibson 1995, Rios and Fanta 1998) and consequently there are limitations in feeding possibilities. Thus, even if attracted to species like G. gibberifrons or L nudifrons, only large specimens ofN. neglecta are able to ingest them. The same was observed for P. antarcticum, which can ingest only small krill (Fanta and Meyer 1998). On the other hand, it was no­ticed that movement, shape, colour and taste were meaningful in active preying, or selection of food.

342 Edith Fanta

N. neglecta N. coriiceps

T. bernacchii

L. nudifrons

P. borchgrevinki

T. newnesi

G. gibberifrons P. antarcticum

B. gigantea

0S W. obesa

Fig. 1. Trophic relations in tanks between the Antarctic fish Notothenia neglecta, N. coriiceps, Trematomus bernacchii, T. newnesi, Pagothenia borchgrevinki, Lepidonotothen nudifrons, Gobio-notothen gibberifrons, Pleuragramtna antarcticum and the krill Euphausia superba, the amphipods Gondogeneia antarctica, Waldeckia obesa and Bovallia gigantea captured during the Antarctic sum­mer in Martel Inlet, Admiralty Bay. Arrows indicate the feeding action of predators towards the preys

and their strength provides an indication of the strength of the trophic relation.

It was observed that some predators were more selective than others. Thus, L. nudifrons, G. gibberifrons, T. newnesi and P. antarcticum fed only on krill, and one species of amphipod, while N. neglecta fed on 7 different preys, including 4 fish species, krill, and two types of amphipods. N. neglecta is known as being ver­satile in its feeding (Kock 1992, Fanta and Meyer 1998, Rios and Fanta 1998), in­gesting whatever is available, and so apparently very well adapted to all possible changes in food availability. It was observed, however, that once preferred food is available, even this species rejects isopods, polychaetes, ophiuroids, and algae (personal observ.).

Fish mostly prefer epibenthos, especially the more motile crustacean groups, to the generally sparse infauna (Daniels, 1982). But it was observed that fish like T. bernacchii, N. neglecta, N. coriiceps and P. borchgrevinki also fed

Laboratory tests on feeding and food preferences of some Antarctic fish 343

on smaller motile fish specimens (Fanta and Meyer 1998). T. bernacchii is an active swimmer that preys on benthic animals (Hureau 1994), but Moreno (1980) also found pelagic euphausiids in their stomachs. In tanks it was seen that they were attracted by krill and amphipods but fed mostly on fish like P. antarcticum, if available. This agrees with Hureau (1970), who states that feed­ing seems to be directed upwards.

Necrophagy seems to be quite common among Antarctic fish (Arnaud 1970). Nevertheless we observed that it occurred only when no motile or live food was available, and only by some of the species, mainly N. neglecta and G. gibberifrons.

It was also observed that fish may be attracted or stimulated either by inade­quate food, rejecting too large individuals or individuals with bad taste or consis­tency or by adequate food items that are well appreciated. Thus, it was noted that some amphipods like G. antarcticum were accepted by all fish except L. nudifrons, being the second most preyed upon of the organisms that were offered, while W. obesa was not identified as food or, when apprehended, was immediately rejected by all species. Krill, on the other hand, was the most preyed upon item and the pref­erential food for most of the fish, stimulating all of them to a high level of activity and feeding behaviour.

Based on all results herein obtained, one can suggest that there are some uni­versal preys like E.superba and G. antarctica, while others are mostly not preyed, like N. neglecta, N. coriiceps, P. borchgrevinki, T. bernacchii and W. obesa. Among the fish, one can conclude that N. neglecta is the strongest predator of this community when in a restricted area while T. newnesi, L. nudifrons, P. antarc­ticum and G. gibberifrons are intensively preyed upon by at least two different predators (Fig. 1).

It may be suggested that stomach content is a useful indicator to determine the presence of certain food items in the natural environment, but not necessarily to de­termine the most appreciated or preferential food. Fish are able to make choices and to reject food that they don't like, even if previously attracted visually or chem­ically towards it. Thus, if there is a great diversity of food available, fish will choose their preferential item. The studies that will follow will test the food prefer­ences from the perspective of the energy costs for feeding and gains from the in­gested nutrients.

Acknowledgements. — The author wishes to thank Dr. Claude De Broyer for giving and identifying the amphipods used as food items; Dr. Vicente Gomes and Dr. Maria Jose" Campos Rocha for the krill and small fish used as food items and for the identification of the krill; to MSc Ana Aparecida Meyer for temporary help in the field work; to the staff of the Brazilian Antarctic Station Comandante Ferraz for valuable help; to the Secretaria da Comissao Interministerial para os Recursos do Mar (SECIRM) and the Brazilian Antarctic Programme for all logistical support. The financial support for die Project was provided by the CNPq grant N25 48131.3/95-8, 480708.96-7 and 480844/97-6; stipends were given to the author by CNPq (N2 521752/95-7 and 300831/93-5). The author is also grateful to Dr. Claude De Broyer and Prof. Dr. Andrzej Kompowski for their valuable suggestions to this paper.

344 Edith Fanta

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Received July 2, 1998 Accepted April 15, 1999

Streszczenie

Praca przedstawia wyniki eksperymentów dotyczących wybiórczości pokarmowej kilku gatun­ków antarktycznych ryb z rodziny Nototheniidae: Notothenia neglecta, N. coriiceps, Trematomus newnesi, T. bernacchii, Lepidonotothen nudifrons, Pagothenia borchgrevinki, Gobionotothen gibbe­rifrons i Pleuragramma antarcticum. W 1000-litrowych zbiornikach w laboratorium umieszczono

346 Edith Fanta

ryby złowione w Zatoce Admiralicji (Wyspa King George, Szetlandy Południowe) i podawano im rozmaity pokarm - ryby, kryl i obunogi. Stwierdzono określoną wybiórczość pokarmową poszcze­gólnych gatunków ryb. Cztery gatunki: N. neglecta, N. coriiceps, P. borchgrevinki i T. bernacchii zjadały rybę Pleuragramma antarcticum. N. neglecta znajdowała się na szczycie piramidy pokarmo­wej, zjadając aż siedem gatunków oferowanych jako pokarm, będąc jednocześnie kanibalem i nekro-fagiem. T. bernacchii i P. borchgrevinki akceptowały 5 gatunków zwierząt podanych jako pokarm. Najbardziej wyspecjalizowaną rybą był T. newnesi, akceptujący jako pokarm tylko dwa gatunki (tab. 1 i 2).

W wyniku tych badań ustalono pewne interakcje troficzne; stwierdzono różnice międzygatun-kowe w preferencjach pokarmowych ryb w warunkach laboratoryjnych przy nielimitowaniu poda­nego pokarmu różnego rodzaju, a także różnice w stosunku do diety obserwowanej w warunkach naturalnych.