Use of wild fish and other aquatic organisms as feed in aquaculture; a review of practices and implications in the Asia Pacific

Global and Asian aquaculture have witnessed a ten-fold increase in production from 1980 to 2004. However, the relative percent contribution to production of each of the major commodities has remained almost unchanged. For example, the contribution of freshwater finfish has declined from 71 to 66 percent in Asia but has remained unchanged globally over the last 20 to 30 years. This fact has dictated trends in the use of fish as a feed for cultured stocks. The growth in the sector has gone hand in hand with an increasing dependence on fish as feed, either directly or indirectly. In a number of countries in the Asia-Pacific region, the aquaculture sector has surpassed the capture fisheries sector in its respective contributions to the gross domestic product (GDP). Aquaculture’s increased contribution to national GDPs can be taken as a clear indication of the contribution of the sector to food security and poverty alleviation. The use of finfish and other aquatic organisms as a feed source can be through direct utilization of whole or chopped raw fish in wet form, through fishmeal and fish oil in formulated feeds, and/or as live fish, although the latter is uncommon and the overall amounts used are relatively small. In the first two categories, the fish used are often termed “trash fish/low-value fish”. Although attempts have been made to define this term, all definitions have a certain degree of ambiguity and/or subjectivity. In this regional review, the amount of fish used as feed sources based on the above categories was estimated primarily from the production data, supported by assumptions on the inclusion levels of fishmeal in formulated feeds and observed feed conversion efficiencies for both formulated feeds and for stock fed trash fish/low-value fish directly. A scenario for the use of fish as feed was developed by starting from the levels of aquaculture production recorded in 2004 and assuming increases in production volumes of 10, 15 and 20 percent by 2010, respectively, for the three trajectories. In parallel, the pattern of wild fish use as feed was projected to change as fish and shrimp farmers increasingly replace farmmade feeds by incorporating trash fish/low-value fish with manufactured feeds that include fishmeal. Also, the fishmeal inclusion rates in manufactured feeds are falling slowly, and this has been incorporated into the projections. The regional review also deals with the production of fishmeal using trash fish/low-value fish in the Asia-Pacific region. Regional fishmeal production as a whole is relatively low when compared with that of major fishmeal-producing countries such as Chile, Iceland and Norway, amounting to approximately 1 million tonnes per year. However, there is a trend towards increasing the use of fish industry waste, such as from the tuna canning industry in Thailand. The fishmeal produced in the region is priced considerably lower than globally traded fishmeal, but its quality is poorer. Total fishmeal use in Asian aquaculture in 2004 was estimated as 2 388 million tonnes, the highest proportion of this being used for crustacean aquaculture (1 418 million tonnes). Based on growth predictions (to year 2010) in the sector and improvements to feed quality and management, it is expected that the quantity of fishmeal used in Asian aquaculture will be slightly less than at present. An estimated 240 000 tonnes of fish oil is used in Asian aquaculture, principally in shrimp feeds. Based on production estimates of commodities in 2004 that rely on trash fish/low-value fish as the main feed source, this regional review suggests that Asian aquaculture currently uses between 2 465 and 3 882 million tonnes, an amount that is predicted to decrease to between 1.890 and 2 795 million tonnes by 2010. The use of trash fish/low-value fish and fishmeal by the aquaculture sector has been repeatedly adjudicated as a non-sustainable practice, and globally the sector is seeking to reduce its dependence on fish as feed through improved feed management practices and development of better quality feeds and feed formulations using alternative ingredients. Over the next few years, decreases in the use of trash fish/low-value fish are also expected to be achieved through better conversion of raw materials into fishmeal and fish oil during the reduction processes. The “way forward” in addressing the issue of the use of fish as feed in aquaculture in the Asia-Pacific region includes the need for a concerted regional research thrust to reduce the use of fish as feed sources in aquaculture, as has been achieved in the animal husbandry sector. Secondly, there is a need to increase farmer awareness on the use of trash fish as feed. This is achievable, considering the similar progress that has been made by the region’s shrimp farming sector, which almost exclusively involves small-scale practitioners who are often clustered in a given locality. The analysis also suggests that the use of trash fish/low-value fish in aquaculture may be compatible with improving food security and alleviating poverty. In Asia, trash fish/low-value fish is mostly landed in areas where there are other suitable fish commodities for human consumption. To make the trash fish/low-value fish suitable and available for human consumption would involve some degree of value-adding and transportation costs, which are likely to increase the price to beyond the means of the consumer, particularly in remote rural areas. Under such a scenario, the direct or indirect use of this perishable resource as a feed source to produce a consumable commodity appears to make economic sense and appears to be the most logical use for overall human benefit. In this manner, trash fish/low-value fish contributes to food security by increasing income generation opportunities and hence contributes to poverty alleviation. Another factor that needs to be taken into account is the large numbers of artisanal fishers who harvest this raw material. The continued use of trash fish/low-value fish, therefore, allows these fishers to maintain their livelihoods¹. Admittedly, this is an area that warrants more detailed investigation, from resource use, livelihoods and economic viewpoints.

Fish oil replacement in starter, grow-out, and finishing feeds for farmed aquatic animals

As aquaculture production continues to grow, there will be an increased use of lipid resources (oils and fats) alternative to fish oil for feed production. The potential for the use of these alternatives varies depending on the feeds in which they are included according to the production phase of the animals to which they are being fed. In starter feeds, where rapid growth, high survival, and normal development are critical priorities, there will remain a need for the use of lipid resources high in omega-3 long-chain polyunsaturated fatty acids (n-3 LC-PUFA). Fish in this starter phase have a critical requirement for the n-3 LC-PUFA docosahexaenoic acid (DHA) and eicosapentaenoic acid (EPA), and fish oils remain the only cost-effective source of these nutrients in the volumes required. However, the greatest demand for lipids is in those diets for the grow-out phase. Most studies on alternative lipid use with animals in this part of the production phase show positive outcomes, in that there are few studies where all the added fish oil cannot be replaced. There are some species, however, where potential replacement levels are suggested to be more conservative, and a general substitution level in this production phase of 75% has been suggested. One of the key effects noted across the grow-out phase is that all alternatives affect the flesh fatty acid characteristics by reducing the level of n-3 LC-PUFA. This issue has provoked the concept of finisher diets, whereby a high n-3 LC-PUFA content diet is fed in order to restore the desired meat fatty acid profiles. Studies examining this concept have found that the tissue triacylglycerol fatty acids were greatly modified and responded in a simple dilution process to the added oil fatty acid composition, whereas the fatty acids of tissue phospholipids were less influenced by dietary fatty acid makeup.

Above- and below-ground vertebrate herbivory may each favour a different subordinate species in an aquatic plant community

At least two distinct trade-offs are thought to facilitate higher diversity in productive plant communities under herbivory. Higher investment in defence and enhanced colonization potential may both correlate with decreased competitive ability in plants. Herbivory may thus promote coexistence of plant species exhibiting divergent life history strategies. How different seasonally tied herbivore assemblages simultaneously affect plant community composition and diversity is, however, largely unknown. Two contrasting types of herbivory can be distinguished in the aquatic vegetation of the shallow lake Lauwersmeer. In summer, predominantly above-ground tissues are eaten, whereas in winter, waterfowl forage on below-ground plant propagules. In a 4-year exclosure study we experimentally separated above-ground herbivory by waterfowl and large fish in summer from below-ground herbivory by Bewick’s swans in winter. We measured the individual and combined effects of both herbivory periods on the composition of the three-species aquatic plant community. Herbivory effect sizes varied considerably from year to year. In 2 years herbivore exclusion in summer reinforced dominance of Potamogeton pectinatus with a concomitant decrease in Potamogeton pusillus, whereas no strong, unequivocal effect was observed in the other 2 years. Winter exclusion, on the other hand, had a negative effect on Zannichellia palustris, but the effect size differed considerably between years. We suggest that the colonization ability of Z. palustris may have enabled this species to be more abundant after reduction of P. pectinatus tuber densities by swans. Evenness decreased due to herbivore exclusion in summer. We conclude that seasonally tied above- and below-ground herbivory may each stimulate different components of a macrophyte community as they each favoured a different subordinate plant species.

Compensatory growth in an aquatic plant mediates exploitative competition between seasonally tied herbivores

The degree to which vertebrate herbivores exploitatively compete for the same food plant may depend on the level of compensatory plant growth. Such compensation is higher when there is reduced density-dependent competition in plants after herbivore damage. Whether there is relief from competition may largely be determined by the life-history stage of plants under herbivory. Such stage-specific compensation may apply to seasonal herbivory on the clonal aquatic plant sago pondweed (Potamogeton pectinatus L.). It winters in sediments of shallow lakes as tubers that are foraged upon by Bewick's Swans (Cygnus columbianus bewickii Yarrell), whereas aboveground biomass in summer is mostly consumed by ducks, coots, and Mute Swans. Here, tuber predation may be compensated due to diminished negative density dependence in the next growth season. However, we expected lower compensation to summer herbivory by waterfowl and fish as density of aboveground biomass in summer is closely related to photosynthetic carbon fixation. In a factorial exclosure study we simultaneously investigated (1) the effect of summer herbivory on aboveground biomass and autumn tuber biomass and (2) the effect of tuber predation in autumn on aboveground biomass and tuber biomass a year later. Summer herbivory strongly influenced belowground tuber biomass in autumn, limiting food availability to Bewick's Swans. In contrast, tuber predation in autumn by Bewick's Swans had a limited and variable effect on P. pectinatus biomass in the following growth season. Whereas relief from negative density dependence largely eliminates effects of belowground herbivory by swans, aboveground herbivory in summer limits both above- and belowground plant biomass. Hence, there was an asymmetry in exploitative competition, with herbivores in summer reducing food availability for belowground herbivores in autumn, but not the other way around.

The role of herbivorous water birds in aquatic systems through interactions with aquatic macrophytes, with special reference to the Bewick’s Swan – Fennel Pondweed system

The role of aquatic macrophytes in stimulating biodiversity and maintaining clear waters is currently undisputed. The management of (eutrophic) shallow waters is therefore often directed at (re-)establishing macrophyte domination. In contrast, the role of water birds has long been considered of minor importance for the functioning of fresh water ecosystems. Indeed, in terms of biomass and production, water birds constitute only a minor part of these systems. However, water birds may graze heavily on water plants under certain circumstances, and the question arises whether herbivorous water birds have an important indirect effect on shallow fresh water systems. Mainly illustrated with the interaction between Bewick’s Swans and Fennel Pondweed, we present data on the role that water plants may play in the life of water birds and how water birds may impact water plants’ fitness in terms of survival, production, dispersal and competitive ability. It appears that water plants may be crucial for water birds during periods of high-energy requirements, such as migration. Despite the plants’ costs associated with water bird grazing, the interaction between water birds and water plants varies in nature from an apparent predator–prey relationship to a mutually beneficial interaction depending on the context and the perspective. For the case of the Bewick’s Swan–Fennel Pondweed interaction, regular bird grazing is sustainable and may actually favour the plant’s dispersal. Thus, Bewick’s Swans themselves may in fact play a crucial role in establishing and maintaining the Fennel Pondweed rich staging sites between the swans’ wintering and breeding grounds, which are vital for the swans’ successful migration.

The effect of seed morphology on the potential dispersal of aquatic macrophytes by the common carp (Cyprinus carpio)

1. The potential for seed dispersal by fish (ichthyochory) will vary among aquatic plants because of differences in seed size and morphology.

2. To examine how seed morphology influences the probability of dispersal by the common carp (Cyprinus carpio), we studied seed ingestion, retention time and subsequent egestion and germination of seeds of Sparganium emersum and Sagittaria sagittifolia, two aquatic plant species with similar sized but morphologically different seeds.

3. We compared dispersal probabilities between the two plant species, in which the probability of dispersal is assumed to be a function of the probabilities of seed ingestion, egestion and germination, and the dispersal distance is assumed to be a function of seed egestion rate over time.

4. We found that, although the soft seeds of S. sagittifolia had an approximately 1.5 times higher probability of being ingested by the carp than the hard seeds of S. emersum (83.15% ± 1.8% versus 56.16% ± 2.7%, respectively), the latter had an almost twofold higher probability of surviving the passage through the digestive tract (38.58% ± 2.7% versus 20.97% ± 1.5%, respectively). Patterns of seed egestion over time did not differ between the two plant species, despite the difference in seed morphology. Gut passage had a different effect on seed germination between plant species. Compared with non-ingested controls, seeds of S. emersum showed a 12.6% increase in germination and a 2.1 day acceleration in germination rate, whereas seeds of S. sagittifolia displayed a 47.3% decrease and 5.1 day delay, respectively.

5. Our results suggest that seed morphology affects the dispersal probability and postdispersal establishment, but not the dispersal distance, of aquatic plants that are dispersed by fish.

Thermal and ventilation performance of a multifunctional sports hall within an aquatic centre

There has been an increasing demand for sports facilities in urban areas recently. As a result of this, more attention is drawn towards not only the energy performance of these building typologies, but also creating a healthy indoor environment for the users. This Study investigates the thermal and ventilation performance of a sports hall within an aquatic centre using computational fluid dynamics (CFD) simulations. IES Virtual Environment software was used to perform the simulations. A number of scenarios were tested by changing the position of extract fans as well as by incorporating natural ventilation strategies. A high level of discomfort was observed in the space. Better comfort condition was achieved by changing the location of exhaust fans ad openings. The results help to recommend some guidelines to inform the proposed refurbishment plans of the site.

Aquatic plant shows flexible avoidance by escape from tuber predation by swans

Deeper burial of bulbs and tubers has been suggested as an escape against below-ground herbivory by vertebrates, but experimental evidence is lacking. As deep propagule burial can incur high costs of emergence after dormancy, burial depth may represent a trade-off between sprouting survival and herbivore avoidance. We tested whether burial depth of subterraneous tubers is a flexible trait in fennel pondweed (Potamogeton pectinatus), facing tuber predation by Bewick's swans (Cygnus columbianus bewickii) in shallow lakes in winter. In a four-year experiment involving eight exclosures, winter herbivory by swans and all vertebrate summer herbivory were excluded in a full-factorial design; we hence controlled for aboveground vertebrate herbivory in summer, possibly influencing tuber depth. Tuber depth was measured each September before swan arrival and each March before tuber sprouting. In accordance with our hypothesis, tuber depth in September decreased after excluding Bewick's swans in comparison to control plots. The summer exclosure showed an increase in tuber biomass and the number of shallow tubers, but not a significant effect on the mean burial depth of tuber mass. Our results suggest that a clonal plant like P. pectinatus can tune the tuber burial depth to predation pressure, either by phenotypic plasticity or genotype sorting, hence exhibiting flexible avoidance by escape. We suggest that a flexible propagule burial depth can be an effective herbivore avoidance strategy, which might be more widespread among tuber forming plant species than previously thought.

Experimental quantification of long distance dispersal potential of aquatic snails in the gut of migratory birds

Many plant seeds and invertebrates can survive passage through the digestive system of birds, which may lead to long distance dispersal (endozoochory) in case of prolonged retention by moving vectors. Endozoochorous dispersal by waterbirds has nowadays been documented for many aquatic plant seeds, algae and dormant life stages of aquatic invertebrates. Anecdotal information indicates that endozoochory is also possible for fully functional, active aquatic organisms, a phenomenon that we here address experimentally using aquatic snails. We fed four species of aquatic snails to mallards (Anas platyrhynchos), and monitored snail retrieval and survival over time. One of the snail species tested was found to survive passage through the digestive tract of mallards as fully functional adults. Hydrobia (Peringia) ulvae survived up to five hours in the digestive tract. This suggests a maximum potential transport distance of up to 300 km may be possible if these snails are taken by flying birds, although the actual dispersal distance greatly depends on additional factors such as the behavior of the vectors. We put forward that more organisms that acquired traits for survival in stochastic environments such as wetlands, but not specifically adapted for endozoochory, may be sufficiently equipped to successfully pass a bird's digestive system. This may be explained by a digestive trade-off in birds, which maximize their net energy intake rate rather than digestive efficiency, since higher efficiency comes with the cost of prolonged retention times and hence reduces food intake. The resulting lower digestive efficiency allows species like aquatic snails, and potentially other fully functional organisms without obvious dispersal adaptations, to be transported internally. Adopting this view, endozoochorous dispersal may be more common than up to now thought.

Space-for-time substitution to assess likely response of aquatic ecosystems to future climate-related change

Predicting and managing ecological response to a changing climate is often limited by an incomplete understanding of response thresholds and biogeographic differences. For example, step changes in rainfall and runoff, and threshold dynamics and hysteresis in ecological response make projection of future conditions difficult. To combat these constraints we propose that biophysical data across exiting climatic gradients can be used in a space-for-time substitution to predict climate-related ecological response elsewhere. This method builds on previous attempts at space-for-time substitution by using patterns in physical and physicochemical data to explain biological differences across the spatial gradient, then using those patterns to formulate hypotheses of temporal ecological response and finally testing those hypotheses on temporal data available in a second, similar region of interest. 

How did this snail get here? Several dispersal vectors inferred for an aquatic invasive species

1. How species reach and persist in isolated habitats remains an open question in many cases, especially for rapidly spreading invasive species. This is particularly true for temporary freshwater ponds, which can be remote and may dry out annually, but may still harbour high biodiversity. Persistence in such habitats depends on recurrent colonisation or species survival capacity, and ponds therefore provide an ideal system to investigate dispersal and connectivity. 2. Here, we test the hypothesis that the wide distributions and invasive potential of aquatic snails is due to their ability to exploit several dispersal vectors in different landscapes. We explored the population structure of Physa acuta (recent synonyms: Haitia acuta, Physella acuta, Pulmonata: Gastropoda), an invasive aquatic snail originating from North America, but established in temporary ponds in Doñana National Park, southern Spain. In this area, snails face land barriers when attempting to colonise other suitable habitat. 3. Genetic analyses using six microsatellite loci from 271 snails in 21 sites indicated that (i) geographically and hydrologically isolated snail populations in the park were genetically similar to a large snail population in rice fields more than 15 km away; (ii) these isolated ponds showed an isolation-by-distance pattern. This pattern broke down, however, for those ponds visited frequently by large mammals such as cattle, deer and wild boar; (iii) snail populations were panmictic in flooded and hydrologically connected rice fields. 4. These results support the notion that aquatic snails disperse readily by direct water connections in the flooded rice fields, can be carried by waterbirds flying between the rice fields and the park and may disperse between ponds within the park by attaching to large mammals. 5. The potential for aquatic snails such as Physa acuta to exploit several dispersal vectors may contribute to their wide distribution on various continents and their success as invasive species. We suggest that the interaction between different dispersal vectors, their relation to specific habitats and consequences at different geographic scales should be considered both when attempting to control invasive freshwater species and when protecting endangered species.

Subfossils of extinct and extant species of Simuliidae (Diptera) from Austral and Cook Islands (Polynesia): anthropogenic extirpation of an aquatic insect?

Subfossil head capsules of Simuliidae larvae have been recovered from swamps on Tubuai and Raivavae of the Austral Islands, and Atiu and Mangaia of the southern Cook Islands. For Tubuai and Raivavae it is likely that the simuliids are extinct, but a single simuliid species is extant on nearby Rurutu. For Atiu and Mangaia, extant simuliids have not been reported, but are known on Rarotonga. Well-preserved head capsules indicate that the Cook Islands subfossils are those of Simulium (Inseliellum) teruamanga Craig and Craig, 1986. For the Austral Islands, the simuliid from Tubuai is considered a variant of Simulium (Inseliellum) rurutuense Craig and Joy, 2000. That from Raivavae is morphologically distinct and is described here as a new species, Simulium (Inseliellum) raivavaense Craig and Porch. Humans arrived in Eastern Polynesia ca. 1,000 years ago resulting in the widespread destruction of lowland forest and conversion of wetlands to agriculture with implied consequences for the indigenous biota of these habitats. Here we consider that one such result was loss of freshwater aquatic biodiversity.