Periphyton ECOLOGY, EXPLOITATION AND. MANAGEMENT Edited by MLE. Azim, M.C.J. Verdegem, \.A\. van Dam and M.C.M. Beveridge 0) 5 a ke BTL ititemole) ONiReye LODO VSR U VY COMME OTA AACA AYE Palen y ME. Azim, Saitama University, Japan. M.CJ. Verdegem, Wageningen University, The Netherlands. \.A. van Dam, UNESGO-IHE Institute for Water Education, Delft, MICA e NGG mer M.C.M. Beveridge, Fisheries Research Services Freshwater Laboratory, Scotland, Uk nD eu Reel al 1 a ne Ue Meco ee ieee tenet en eset ‘on submerged substrates, including associated non-attached organisms and detritus. Thus the Pal ab etc eon a oer Cor pI see ae ote) on ee CNS orgs CO clon tole Mareloamcouen tela eter ea | fixation and nutrient cycling in aquatic ecosystems; as an important source of food in aquatic t Bea Se WCC ui le-tcome Ken inonenie epee Deere Wa ae ee a Demo sacs seo ees Ue note ku ace arecticUh eto oer ea exploitation part covers its nutritive qualities and utilization by organisms, particularly in Se re mL UE Re Ra Nts Noll ah mer eae eta tere) and its effects on water quality and animal health in culture systems. The first Cou oo Sonne aaa oe eae eee ECTS oes erie ts NEE ieee an sient Bacteria from Fish and Other Aquatic Animals: a Practical Identification Manual Aquaculture: an Introductory Text Aquaculture and Fish: Biotechnology: Genetic Approaches Verdegem van Dam BeveridgeForeword Contents About the Editors Contributors Chapter 1 Periphyton and Aquatic Production: an Introduction ME. Azim, 11. 12. 13. 14, 15. 16. 17. ‘M.C.M. Beveridge, A.A. van Dam and M.C.J. Verdegem Definition and Synonyms Contribution of Periphyion to Aquatic Productivity Status of World Aquaculture: Systems, Production and Trends Aguacultural Ecology ‘The Role of Periphyton in Aquaculture Ponds 1.5.1. Phytoplankton-based food web 1.5.2. Periphyton-based food web: the periphyton loop 1.5.3. Detritus-based food web 1.5.4. Fish yields Periphyton-based Aquaculture: an Overview Outline/Structure of the Book References Part I EcoLocy Chapter 2 Periphyton Structure, Diversity and Colonization M.E. Azim and T. Asaeda 24 22. 2.3. 2.4. 2.5. 2.6. Introduction Morphological Structure Biochemical Composition Taxonomic Composition and Diversity Colonization Process” Sloughing and Recolonization ‘Acknowledgements References bas Bawnaanceanne = 15 15 15 7 19 23 27 29 29 vilvill Contents Chapter 3. Periphyton Dynamics and Influencing Factors AE. Vermaat 3.1. Introduction 3.2. Patterns in Time: Colonization, Attachment, Growth, Dislodgement and Seasonality 3.3. Constituents of Periphyton Mass 3.4. Primary Abiotic Factors 3.4.1. Light and temperature 3.4.2. Nutrients 3.4.3. Dissolved inorganic carbon 3.5. Biotic Interactions 3.5.1. Substrate specificity: host-periphyton interactions 3.5.2. Grazing 3.6. Patterns in Space: From mm to km Scale 3.7. Modelling Periphyton Dynamics References Chapter 4 Periphyton in the Aquatic Ecosystem and Food Webs RG, Wetzel 4.1. Periphyton as a Critical Metabolic Component of Aquatic Ecosystems 4.2. Fluxes of Organic Matter and Eneray in Aquatic Ecosystems 4.3. Structure and Couplings among Periphytic Organisms 4.4. Productivity of Periphyton Communities: Growth Regulation and Habitat Variables 4.8. Recycling: the Key to Nutrient Fluxes and Maximizing Productivity of Periphyton 4.6. Modulation of Periphyton by Light Availabilty 4.7. Modulation of Periphyton by Mortality and Losses - What Do ‘They Mean to Higher Trophic Levels? 4.8. Potential Effects of Global Changes in Climate and Related Environmental Conditions References Chapter 5 Periphyton in Freshwater Lakes and Wetlands L.G. Goldsborough, R.L. McDougal and A.K. North 5.1. Introduction 5.2. Primary Production 5.3. Nutrients 5.4. Light 5.5, Hydrodynamics 5.6. Herbivory 5.7. Periphyton-Macrophyte Interactions 5.8. Periphyton in Lentic Freshwater Food Webs 5.9. Conclusions Acknowledgements References 35 35 36 37 40 40 41 42 43 43, 45 51 SL 52 53 55. 63 65 66 71 71 73 75 77 78 78 79 81 82 82 83Contents Part II EXPLOITATION Chapter 6 Utilization of Periphyton for Fish Production in Ponds: a Systems Ecology Perspective A.A. van Dai 61. 6.2. 6.3. 64. 65. 6.6. 67. m and M. Introduction Experimental Reference: Bangladesh Model 1: Phytoplankton-based Fish Production in Ponds Model 2: Adding the Periphyton Loop Model Construction and Implementation Results Discussion Verdegem References Chapter 7 Adaptations to Feeding in Herbivorous Fish (Cyprinidae and Cichlidae) FA. Sibbing Th. 72 73, 74. 1a 76. Ua 78. 29, 7.10. and F. Witte Introduction 7.1L. Plants as food 7.1.2. Herbivorous fish Feeding Modes for Cropping Periphyton ‘Structure and Function of the Feeding Apparatus in Fish 7.3.1. The fish head as construction for feeding Feeding Mechanisms in Herbivorous Fish Foraging on Algae and Plants 7.5.1. Search and detection 7.5.2. Intake or cropping Processing Algae and Plants 7.6.1. Selection and retention 7.6.2. Food transport towards chewing cavity and oesophagus 7.6.3. Pharyngeal mastication 7.6.4. Digestion Allometry in Feeding Structures and Diet Shifts Periphyton Feeders among Cyprinids Periphyton Feeders among Cichlids Conclusions and Perspectives Acknowledgements References Chapter 8 Traditional Brush Park Fisheries in Natural Waters RL. Welcomme 8.1. 8.2. as Introduction ‘Nature of Brush Parks 8.2.1. Construction 8.2.1.1. Form 8.2.1.2. Types of wood and vegetation used 8.2.2. Location 8.2.3. Distribution 8.2.4. Area Ecology of Brush Parks 8.3.1. Trophic Status 8.3.1.1, Periphyton 91 o1 92 92 95 97 102 109 113 113 113 115 115 116 116 7 118 118 118 121 12. 125 125 130 131 132 132 134 135 135 141 141 141 141 141 142 144 144 144 144 144 144x Contents 8.3.12. Decay 146 8.3.1.3. Associated organisms 146 8.3.2. Reproduction 146 8.4. Operation 146 8.4.1. Supply of food 146 8.4.2. Density of planting 146 8.4.3. Fishing 147 8.4.3.1. Method 147 8.4.3.2. Frequency 148 8.4.3.3. Species caught 148 8.4.3.4. Yield 148 8.5. Management 148 8.5.1. Feeding 148 8.5.2. Frequency of fishing 150 8.5.3. Social and economic aspects 151 8.6. Environmental Impacts 152 8.6.1. Removal of fish 152 8.6.2. Deforestation 152 8.6.3. Siltation 153 8.7. Models of Management 153 8.7.1. Parks with non-breeding fish populations 153 8.7.2. Parks with breeding populations 153 8.8. Evaluation 154 8.9. Management Strategies 155 8.9.1. Excessive removal of fish and risks of parks contributing to overfishing 155 8.9.2. Conflict between gears 155 8.9.3. Deforestation 155 8.9.4, Siltation 155 8.9.5. Social problems 155 8.9.6. Financial 156 ‘Acknowledgements 156 References 156 Chapter 9 Periphyton as Biological Indicators in Managed Aquatic Ecosystems 159 S. Sabater and W. Admiral 9.1. Introduction 159 9.2. Periphyton as Receptors of Physical, Chemical and Biological Disturbances 160 9.2.1. Effects of toxicants and occurrence of nuisance organisms 161 9.2.2. Light attenuation and excess 162 9.3, Structural and Functional Responses of Periphyton to Disturbance 162 9.4. Using Periphyton for Monitoring Managed Aquatic Systems 166 9.4.1, Use of pigments as monitoring tools 166 9.4.2. Periphyton/biofilm growth forms. 167 9.4.3. Species consortia 168 9.4.4. Monitoring the productivity of photosynthetic periphyton 170 Acknowledgements im References 171Contents xi Part Il MANAGEMENT Chapter 10 Effect of Periphyton on Water Quality 179 A. Milstein 10.1. Introduction 179 10.2. Water Quality in Periphyton-based Aquaculture Ponds 180 10.2.1. Periphyton-environment relationships in the absence of grazing pressure by cultured organisms 181 10.2.2. Periphyton-environment-cultured organisms relationships 183 References 188 Chapter 11 Similarities between Microbial and Periphytic Biofilms in Aquaculture Systems 191 M.CuJ. Verdegem, E.H. Eding, V. Sereti, R.N. Munubi, R.A. Santacruz-Reyes and A.A. van Dam 11.1. Introduction 191 11.2. Structure of Biofilms and Periphyton Mats 192 11.3. Ammonia Removal 192 11.3.1. Biofilms 192 11.3.2. Periphyton mats 193 114. Solid Removal and Detachment 197 11.5. Oxygen Consumption 198 11.5.1. Biofilms 198 11.5.2. Periphyton mats 199 11.6. Biofilm and Periphyton-based Recirculating Aquaculture Systems 200 11.6.1. The concept of algal turf scrubbers 200 11.6.2. Periphyton-based recirculating aquaculture systems 201 References, 202 Chapter 12 Periphyton-based Pond Polyculture 207 ME. Azim and M.A. Wahab 12.1. Introduction 207 12.2. Polyculture System 207 12.3. Suitable Fish Species for Polyculture 208 12.4. History of Polyculture 208 12.5. Development of Periphyton-based Polyculture Technology 210 12.5.1. Indian major carp polyculture 210 12.5.1.1. Selection of fish species by monoculture trials 210 12.5.1.2. Tworspecies polyculture 212 12.5.1.3. Three-species polyculture 213 12.5.2. Tilapiafreshwater prawn polyculture 214 12.6. Intensification of Periphyton-based Polyculture 214 12.7. Possible Mechanisms for Enhanced Fish Production in Periphyton-based Systems 215 12.7.1. Provision of additional food and shelter 215 12.7.2. Opportunity for higher nutrient efficiency 215 12.7.3. Better environment through improved water quality 216 12.7.4. Optimize the synergistic effects 217 12.8. Applicability of the Periphyton Technology and its Socio-economic Impacts 218 Acknowledgements 218 References 219xii Contents Chapter 13 Research on Periphyton-based Aquaculture in India. 223 P. Keshavanath and B. Gangadhar 13.1. Introduction 223 13.2. Periphyton and Plankton Productivity 224 13.3. Fish Production 225 13.3.1. Substrate evaluation studies 225 13.3.2. Studies with substrates and supplemental feed 227 13.3.3. Stocking density and production 231 13.3.4. Use of substrates in nursery rearing 232 13.4. Biofilm in Fish Health, 233 13.5. Water Quality in Substrate-based Systems 233 13.6. Digestibility of Periphyton 233 13.7. Economics of Substrate-based Fish Culture 234 13.8. Conclusions 235 References 235 Chapter 14 Periphyton-based Cage Aquaculture 237 SMH. Huchette and M.C.M. Beveridge 14.1, Cage Aquaculture — an Overview 237 14.2. Biofouling 238 14.3. Periphyton-based Cage Aquaculture 239 14.4. Discussion 240 References 244 Chapter 15 Utility of Added Substrates in Shrimp Culture 247 J.H. Tidwell and D. Bratvold 15.1. Introduction 207 15.2. Marine Shrimp 247 15.2.1. Marine shrimp nursery systems 248 15.2.2, Marine shrimp juvenile culture systems 249 15.2.3. Conclusions 254 15.3. Freshwater Prawns 254 15.3.1. Natural history 255 15.3.2. Importance of natural foods 255 15.3.3. Concept of added substrate 260 15.3.4. Recent developments 261 15.3.5. Conclusions 265 References 265 Chapter 16 Importance of Periphyton in Abalone Culture 269 T. Kawamura, B.D. Roberts and H. Takami 16.1. Introduction 269 16.2. Life History of Abalone 270 16.3. Growth Forms and Cornmunity Succession in Marine Benthic Diatoms 270 16.4. Abalone Larval Settlement 272 16.4.1. Effects of diatom films (periphyton) on the settlement of larval abalone 272 16.4.2. Other settlement methods 273 16.4.2.1. Pre-grazed plates 273 16.4.2.2. Ulvella lens 273 16.4.2.3. -Aminobutyric acid (GABA) 273Contents 16.4.2.4. Bacteria 16.4.2.5, Crustose coralline algae (CCA) 16.5. Nutrition of Young Abalone 16.5.1. Phases of nutrition in abalone 16.5.1.1. Phase 1: larval development 16.5.1.2. Phase 2: initial post-arval development 16.5.1.3, Phase 3: growth of postlarvae from ~400 to ~700 um SL 16.5.1.3.1. Factors controlling diatom ingestion by small abalone 16.5.1.4, Phase 4: postlarvae from 0.7 mm to several millimetres SL 16.5.1.5. Phase 5: transition to macroalgal or manufactured diets 16.5.2. Other food sources 16.6. Practical Application of Periphyton to Abalone Culture 16.6.1. Key diatom species for abalone hatcheries 16.6.2. The effectiveness and limitations of the pre-grazed plate method 16.6.3. Use of GABA with diatoms for triggering settlement 16.6.4. Other food sources 16.7. Concluding Remarks References Chapter 17 Periphyton-based Aquaculture in Asia: Livelihoods and Sustainability S.W. Bunting, M, Karim and M.A, Wahab 17.1. Introduction 17.2. Research Outcomes 17.2.1. Knowledge and assets influencing uptake 17.2.2. Putting vulnerability in context 17.2.3. Conflicts associated with uptake 17.2.4. Environmental impacts 17.2.5. Livelihood outcomes 17.26. Broader issues influencing uptake and adoption 17.2.6.1. Attributing outcomes to periphyton-based aquaculture 17.2.6.2. Substrate selection 17.2.6.3. Substrate management 17.2.6.4. Social capital 17.3. Conclusions Acknowledgement References Chapter 18 Periphyton Ecology, Exploitation and Management: Knowledge Gaps and Directions for Future Research M.CJ. Verdegem, A.A. van Dam, M.E, Azim and M.C.M. Beveridge 18.1, Introduction 18.2. Ecology 18.2.1. The role of periphyton in natural systems 18.2.2. Periphyton structure and new monitoring techniques 18.2.3, Optimal grazing pressure 274 274 274 274 274 274 275 275 276 276 277 278 278 278 280 280 280 280 285 286 287 289 289 290 290 291 291 292 293 295 295 296 297 299 299 299 299 300 301Contents, 18.3. Exploitation 18.3.1. Abilities to harvest and assimilate periphyton 18.3.2. Nutritive quality of periphyton 18.3.3. Regeneration capacity of periphyton under grazing pressure 18.4. Management 18.4.1. Increased production from periphyton-based production studies 18.4.2. Polyculture 18.4.3. Water quality . Health and well-being .Disseminating periphyton technology Periphyton-based aquaculture: an ecological approach 301 301 302 302 303 303 304 304 305 307 307 3111 Periphyton and Aquatic Production: an Introduction M.E. Azim,! M.C.M. Beveridge,? A.A. van Dam? and M.C,J. Verdegem* 'Graduate School of Science and Engineering, Saitama University, Japan; 2ERS Freshwater Laboratory, Pitlochry, UK; “Department of Environmental Resources, UNESCO-IHE Institute for Water Education, The Netherlands; “Aquaculture and Fisheries Group, Department of Animal Sciences, Wageningen University, The Netherlands. 1.1. Definition and Synonyms Periphyton is of growing interest worldwide. Itis immensely important for various reasons: (i) it is a significant, and often the dominant, contributor to carbon fixation and nutrient cycling in aquatic ecosystems; (i) it is an excel- lent indicator for changes occurring in the aquatic environment; (ii) it is used to improve water quality in lakes and reservoirs; (jv) it increases food availability in aquatic production systems; (v) it can provide specialty foods for fish and shellfish larvae culture; and (vi) it can be used for wastewater treatment. The variety of organisms found in periphyton is enormous; it harbours algae, bacteria, fungi, protozoa, z00- plankton and other invertebrates. These organ- isms function as a community, making them highly efficient in capturing and processing nutrients. The term ‘periphyton’ in classical limnol- ogy refers to the microfloral community living altached to the surfaces of suimmerged objects in water (Wetzel, 1983a). This definition does not include fungal, bacterial, protozoan and other attached animal components, which are included in the German word ‘aufwuchs’ ‘Aufwuchs includes all the organisms that are attached to, or move upon, a submerged sub- strate, but which do not penetrate into it, ‘whereas periphyton refers to the total assem- blage of sessile or attached organisms on any substrate (Reid and Wood, 1976; Weitzel, 1979}. The difference is in the unattached organisms that are often found in association with the periphyton assemblage. Sometimes, the terms ‘euperiphyton’ (immobile organisms attached to the substrate by means of rhizoids, gelatinous stalks or other mechanisms) and ‘pseudoperiphyton’ or ‘metaphyton'’ (free-liv- ing, mobile forms that creep among or within the periphyton) are used (Weitzel, 1979). The term ‘biofilm’ is preferred in other fields of application, such as wastewater treatment (Cohen, 2001), drinking water technology (Momba et al., 2000), food processing (Joseph et al., 2001) and dentistry (Rosan and Lamont, 2000), where it is used mainly with reference to attached bacteria and protozoa but not algae {O'Toole et al., 2000). Some authors prefer to call them ‘attached algae’ or ‘attached microor- ganisms’, but these terms disregard the many other forms that live in periphyton assem- blages. Periphyton also has different synonyms depending on the substrate on which it grows {Table 1.1). Throughout this book, the term periphyton is used to refer to the entire complex ‘SCAB International 2008. Perjphyton: Ecology, Eyploitation and Management (eds M.E. Azim eta.) 12 ME. Azim etal. Table 1.1. Category of periphyton based on the types of substrates. Substrates ‘Synonyms Aquatic plant Epiphyton Sedimentimud/sitt Epipelon ‘Wood Epixylon Rook Epilthon Sand Epipsammon of attached aquatic biota on submerged substrates, including associated non-attached ‘organisms and detritus (van Dam et al., 2002). Thus, the periphyton community comprises bacteria, fungi, protozoa, phytoplankton, z00- plankton, benthic organisms, organic detritus and a range of other invertebrates and their larvae, 1.2. Contribution of Periphyton to Aquatic Productivity Periphyton is immensely important in aquatic systems as it provides community structure and primary productivity that support a wide range of aquatic organisms. Periphyton is easily grazed upon by small invertebrates, fishes and shrimps and hence contributes considerably to the productivity of aquatic ecosystems, natural or human-made. In many water bodies, the contribution of the periphyton community to production is greater than that of the phytoplankton. For example, in a study that compared primary pro- duction in a turbid and clear lake, phytoplank- ton was found to account for 96% of the total annual production in the turbid lake while epipelon contributed 77% in the clear lake (Liboriussen and Jeppesen, 2003). However, total annual production of phytoplankton and epipelon was 34% greater in the turbid lake (190 g C/m?/year) than in the clear water lake (141 g C/m?/year). In some cases, the contribution of periphyton to annual primary productivity can be as high as 1 kg C/m? (Goldsborough and Robinson, 1996). In some earlier studies, it was shown that periphyton can have an important share (42% for the lake studied) of the total annual production, espe- cially in shallow lakes with large littoral zones. (Wetzel, 1964). In five oligotrophic lakes, for example, periphyton contributed 43-97% of the total productivity in the shallow (2-3 m) zone (Loeb et al., 1983). In general, high values for periphyton pro- ductivity are found on coral reef systems, with values typically in the range of 1-3 g C/m?/day (Carpenter, 1986; Polunin, 1988; Klumpp and Polunin, 1989; van Rooij et al., 1998). Guiral et al. (1993) reported the highest value for peri- phyton production of 7.9 g C/m?/day in acadja systems (artificial brush parks; see Welcomme, Chapier 8 this volume), which was 4.5 times higher than that of the lagoon phytoplankton community, The authors also reported that 80% of the acadja productivity was due to periphyton. In ponds provided with periphyton substrates, half of the primary productivity was contributed by periphyton (1.7 g C/m?/day) (Azim et al., 2002). However, the contribution of periphyton on the pond bottom. and on aquatic vegetation was excluded from the calculation, indicating that the total peri- phyton productivity is likely to be higher than that of phytoplankton in ponds provided with artificial hard substrates 1.3. Status of World Aquaculture: Systems, Production and Trends Fish production through aquaculture is realized ina wide variety of culture systems, depending on farm location, the nature of the water envi- ronment and culture intensity. Aquaculture sys- teins may be land-based or water-based. Land-based systems are built on dry land and include mainly ponds, raceways and rice fields Water-based systems are usually situated inland or in sheltered coastal waters, and include enclosures, cages, pens and rafts. The majority of aquaculture production comes from land- based freshwater culture systems. Based on the ‘water environment, aquaculture systems can be divided into freshwater, brackish water and marine systems. ‘Aquaculture systems may be categorized as extensive, semi-intensive and _ intensive, although there are no rigid boundaries among categories. Production in extensive pondPeriphyton and Aquatic Production 3 systems is based on the natural productivity of the pond and solar energy. It involves traditional practices relying on natural food and is charac- terized by low fish production. Often the life cycle is not completely controlled. An example is extensively operated fish ponds that rely on the supply of wild juveniles and are pro- vided with feed and fertilizers only occasionally In semi-intensive systems, fish are purposely stocked and the natural food supply is enhanced through the use of organic and inorganic fertiliz~ ers and low-cost supplemental feed such as rice bran, oil cake and other agricultural by-prod- ucts. Today, extensive and semi-intensive aqua culture systems contribute the bulk of the ‘world’s aquaculture production. Intensive systems are based predominantly on high-quality complete feeds and produce a greater output per unit production system. This is achieved through higher levels of tech- nology and a greater degree of management. “The cultured species are often reared fram egg to adult stage within the culture facilities, and stocked in smaller, well-designed facilities at higher densities. Water quality is carefully main- tained with the aid of biofilters, solid removal units, pumps, aerators and automated water quality recording systems. According to the Food and Agriculture Organization (FAO) statistics, the contribution of aquaculture to global supplies of fish, crus- taceans and molluscs continues to grow, increasing from 3.9% of total production by weight in 1970 to 27.3% in 2000. Aquaculture is the fastest growing animal production sector. Worldwide, the sector increased at an average compounded rate of 9.2% annually since 1970, compared with only 1.4% for capture fisheries and 2.8% for lanc-based meat production sys- tems. In 2000, the reported total aquaculture production (including aquatic plants) was 45.7 million tonnes by weight and US$56.5 billion by value, Production is dominated by Asian coun- tries, particularly China, India, Indonesia, Bangladesh, Thailand and Vietnam, which together account for 90% of world aquaculture production. Freshwater finfish, especially Chinese and Indian carp species, accounts for the greatest share (52%) of total aquaculture production. It is particularly significant that aquaculture production in developing countries and low-income food-deficit countries {LIFDCs) has been growing steadily at an aver- age rate of about 10% per year since 1970. By contrast, aquaculture production within devel- oped countries has been growing at an average rate of only 3.7% per year since 1970, and even showed a decrease of 2.4% from 1999 to 2000. With the exception of marine shrimp, in 2000 the bulk of aquaculture production in developing countries comprised omnivorous or herbivorous fish or filter-feeding species cultured mainly in semi-intensive ponds. In contrast, 74% of finfish culture production in developed countries were carnivorous species relying on artificial complete diets (FAO, 2002). During the past three decades, aquaculture has expanded, diversified, intensified and made technological advances. The potential of this development to enhance local food security, alleviate poverty and improve rural liveli- hoods has been well recognized. The Bangkok Declaration and Strategy (NACA/FAO, 2000) emphasizes that the practice of aquaculture should be pursued as an integral component of development, contributing towards sustainable livelihoods for poor sectors of the community, promoting human development and enhancing social well-being. Aquaculture is not only seen as an important contributor to development, but also as an important tool for increasing food security, especially in the developing world. Rural aquaculture, including enhanced fisheries and cullure-based fisheries, has made signifi cant contributions to the alleviation of poverty, This has been achieved directly through small- scale farming households raising aquatic organ- isms for domestic consumption and/or income generation. Indirectly, it has provided employ- ment and supplied low-cost fish to local mar- kets. In Asia, where most fish farmers are relatively poor, there is vast potential for the poorer section of the society to become involved in aquaculture. There are wide oppor- tunities for the poor to integrate aquaculture into existing farming systems and thus there is still room for growth in aquaculture production (Lewis, 1997; FAO, 2000). Unfortunately, efforts to develop low-cost rural aquaculture technologies so far have not led to large-scale increases in fish production from small inte- grated farming systems. Because the majority of the farmers in Asia are poor, many cannot provide supplemental4 ME. Azimetal. feed to their ponds. Feed-driven systems, how: ever, are also inefficient. The conversion effi- Gieney of feed isaround 20% at best (Davenport et al., 2003), the remainder being lost to the environment as wastes, of which only a part is subsequently recycled within the systems. The sustainability of intensive feed-driven, especially fishmeal- and fish-oil-based, aquaculture is questionable (Naylor et al., 2000). From eco- nomic and environmental points of view, there is a need fo examine options to make aquacul- ture systems more resource-efficient. The big challenge for scientists, then, is to increase fish production to meet increased demands by an ‘expanding human population without jeopard- izing the environment. 1.4. Aquacultural Ecology In aquaculture ponds, complex interrelated physical, chemical and biological processes contribute to the formation and the stability of the ecosystem. For simplicity, three basic ecological processes can be distinguished: production, consumption and decomposition (Fig. 1.1). Production is based on the conver- sion of solar energy, carbon dioxide and dis- solved inorganic nutrients into plant biomass and oxygen through photosynthests. It eads to the formation of phytoplankton, periphyton and other submerged plants, providing the base of the aquatic food web. This base can be sup- plemenied by the addition of organic matter in the form of manure or artificial feed. In the consumption process, both autochthonous and added organic matters are eaten directly or indi- rectly by aquatic animals such as zooplankton, Fig, 141. Aquacultural ecology: nutrient exchange through production, consumption and decomposition processes. benthos, fish or shrimp, providing energy and building blocks for biomass. Decomposition is the third basic ecological process, consisting of the breakdown and mineralization by microor- ganisms (bacteria, protozoa, fungi) of organic matier, leading to the formation of detritus and inorganic nutrients. The detritus is consumed directly by the cultured fishes or crustaceans or by other small animals on which the cultured species feed (Colman and Edwards, 1987; Moriarty, 1997). The inorganic nutrients are recycled to stimulate primary production and serve as the base of autotrophic food webs. So, ina practical sense, there is no clear distinction between the sources of nutrients used by the animals under culture. Nutrients are transferred or exchanged between different trophic levels through these ecological processes, which are controlled by a range of biotic and abiotic fac- tors (Fig. 1.1), Whether primary production (phytoplank- ton) can be harvested directly by fish depends largely on the fish species stocked. Because many fish, especially carnivorous species, are not able to harvest phytoplankton directly from the water column, one or more extra trophic levels are involved in converting phytoplankton into fish biomass. With an estimated energy transfer efficiency of 10% per trophic level (Pauly and Christensen, 1995), maximum fish yields may not be more than 1% of the eneray fixed by autotrophs. As a result, fish yields from extensive and semi-intensive ponds could be increased by a factor of up to ten if fish could exploit the primary production directly. Fish species that feed lower in the food chain, i.e herbivorous and omnivorous fish, are the most efficient to culture in earthen ponds, although ee "| Pa \ Natens cor P Decomposition ‘eatenPeriphyton and Aquatic Production 5 such species do not always realize the highest market prices. 1.5. The Role of Periphyton in ‘Aquaculture Ponds ‘A common assumption has been that the phy- toplankton community is the most important component in the aquatic food web, particularly {in aquaculture ponds, in terms of energy fixa- tion and fuelling of the food web. Therefore, in. traditional aquaculture, strategies adopted for reducing inputs and increasing efficiencies of aquaculture production have focused mainly on optimization of exploitation of the phytoplank- ton-based food web. The periphyton-based food web offers possibilities for further optimiz- ing input utilization in extensive and semi-inten- sive aquaculture ponds. Fishes and crustaceans can also exploit the detritus-based food web. In subsequent sections, these three basic food ‘webs are discussed, giving special attention to periphyton-based food webs. Figure 1.2 offers a preliminary conceptual model of the three food webs and their interactions. 1.5.1. Phytoplankton-based food web Besides being a food source, phytoplankton serves a number of very important functions in pond aquaculture (see section 1.4). Itis a net producer of dissolved oxygen, which is indis- pensable for fish growth and production (Smith, and Piedrahita, 1988; Teichert-Coddington and Green, 1993). Its also the most important sink of ammonia-nitrogen, which is excreted by fish and is potentially toxic (Hargreaves, 1998; Jiménez-Montealegre, 2001). Nevertheless, there are some disadvantages in phytoplank- ton-based production. Phytoplankton in ponds often forms dense blooms, resulting in large diurnal variations in dissolved oxygen con- centrations, from supersaturation during the day (due to photosynthesis) to oxygen deple- ‘ion in the early morning (due to night-time res- piration). Blooms can be unstable and crash, ‘decompesiion Fig. 1.2. Preliminary conceptual model ofa periphyton-based aquaculture system. Only major rate and state variables are shown. The shaded area indicates the ‘periphyton loop’.6 ME. Azim etal. resulting in oxygen depletion and fish mortality (Madenjian et al., 1987; Delincé, 1992) Decomposition of settled phytoplankton at the pond bottom may result in toxic decomposition products (ammonia, nitrite, methane, hydrogen sulphide) and increase sediment oxygen demand. 1.5.2. Periphyton-based food web: the periphyton loop To date, most research on periphyton has been conducted in lakes, rivers, wetlands and coral reefs, focusing on biomass and productivity and their determining factors, taxonomic composi- tion, and periphyton sampling methods. There is litle information on periphyton-based food webs, especially in managed ecosystems such as ponds, Most herbivorous carp and tilapia species cannot meet their full energy demand by filter- feeding on planktonic algae only (Dempster et al., 1995). Besides phytoplankton, herbivo- rous fish generally require larger-sized food sources such as benthic algae (periphyton), algal-based detritus or higher aquatic plants that can be harvested more efficiently (Dempster et al., 1993; Yakupitiyage, 1993). Benthic algal mats rarely grow on the bottom of highly eutrophic ponds due to light limitation. To grow, they need some hard substrate within the pond’s euphotic zone, which is scarce in tradi- tional fish ponds. Any materials that provide surface area can be used in aquaculture ponds to support periphyton production. The addi- tional shelter provided by the substrate allows more of the resources to flow into fish biomass (Fig, 1.2). The new primary production and benthic secondary production of the attached communities fostered by the artificial substrate supports a new food web, part of which ends up as fish biomass (Miller and Falace, 2000), Grazing on a two-dimensional layer of periphy- ton is mechanically more efficient than filtering algae from a three-dimensional planktonic envi- ronment (Dempster et al., 1993; Horne and Goldman, 1994). Most truly herbivorous fish species feed on larger, benthic, epilithic or peri- phytic algae, rather than on phytoplankton (Horn, 1989). Aquaculture production in ponds provided with periphyton substrates is higher than in ponds without substrates (Azim, 2001; van Dam et al., 2002), If pond algae could be grown on sub- strates, more fish species would be able to harvest them, resulting in a more efficient uti- lization of primary production. Communities of attached algae are generally more stable than phytoplankton and the risk of collapse is lower {Westlake et al., 1980). Some studies suggest that the production of attached algae per unit water surface area is higher than that of phyto- plankton (Wetzel, 1964}. However, periphyton biomass and productivity might be influenced by both ecological (temperature, sunlight, etc.) and operational processes (fish stocking density or grazing pressure, fertilization, substrate den- sily and type, etc.). Grazing of periphyton by fish or invertebrate grazers can reduce biomass, but at the same time increase the productivity of periphyton (Hatcher, 1983; Hay, 1991) Because grazing removes dead or old algae and enhances light and nutrient availabilty to the ‘mat, the algal assemblages are maintained in a more healthy, productive and early sueces- sional stage (Huchette et al., 2000). When substrates are installed in ponds, the flux of nutrients through the periphyton ‘loop’ as well as the phytoplankton ‘loop’ is increased ig. 1.2). In consequence, the overall nutrient efficiency of the system increases. Periphyton production effectively doubled autotrophic car- bon production (on average, 1.2 g C/m?/day from periphyton and 1.3 g C/m?/day from phytoplankton) (Azim et al., 2002). However, sometimes a trade-off between phytoplankton and periphyton occurs, especially in highly fer- tilized ponds (Azim et al., 2001), but this can be minimized by stocking fish species with comple~ mentary feeding niches (Azim and Wahab, Chapter 12 this volume) 1.5.3. Detritus-based food web Detritus refers to non-living organic matter, which includes both particulate and dissolved matter. It is an important food source in aqua- culture ponds (Bowen, 1987; Schroeder, 1987; Fig. 1.2). Anaerobic processes are sig- nificant in the decomposition of detritus inPeriphyton and Aquatic Production 7 pond sediments, where oxygen may be fully depleted within the top 1 mm, due to the very rapid bacterial growth and poor diffusion of ‘oxygen (Moriarty, 1997). The estimated contribution of phytoplank- ton to sediment organic matter accumulation in the pond bottom ranges from 20 to 50% of phytoplankton standing stock per day (Jiménez-Montealegre, 2001). Thus, a large part of phytoplankton production is decom- posed at the pond bottom and contributes to the accumulation of nutrients in the sediment. Jn contrast, after dying, periphytic cells remain in place, providing a rich source of organic nutrients for the heterotrophic organisms asso- ciated with the periphyton layer. Processing of this organic matter yields inorganic nutrients that can be utilized by the living algae again (Wetzel, 1983b). On the other hand, if the graz- ing pressure is insufficient in relation to peri phyton production, periphyton mats may be dislodged from the substrate, thereby contribut- ing to the turbidity in the water column (Milstein et al., 2003). 1.5.4, Fish yields In a periphyton-based fish pond, fish yields depend on periphyton substrate density, quan- tity and the nutritional quality of the periphyton, the grazing efficiency of the cultured fish species, fish species combination and density, etc. Fig. 1.2). In polyculture ponds, using bam- boo as substrates at submerged surface area densities equivalent to 50, 75 and 100% of the pond water surface area, fish production was found to increase linearly with increasing substrate surface area (Azim et al., 2004). No ‘maximum production was reached within the range of densities used. The nutritional quality of periphyton is highly variable, depending particularly on habi- tats and substrate types and grazing pressure. The overall nutritional quality of periphyton is much better in closed waters, especially in aquaculture ponds, than in open waters (see Azim and Asaeda, Chapter 2 this volume; van Dam et al., 2002). This is mainly because peri- phyton is purposely grown in managed ponds by positioning substrates in euphotic zones, However, it is very important that the nutri tional quality of periphyton is sufficient to sup- port the dietary needs of cultured animals; otherwise, supplemental feed should be applied to the ponds. Not all fish are able to graze on periphyton; morphological and physiological adaptations to periphyton grazing are required (Sibbing and Witte, Chapter 7 this volume), However, more insight is still needed into the mechanisms for periphyton exploitation through selective feeding by different fish com- munities. Periphyton-based Aquaculture: an Overview ‘The idea of periphyton-based aquaculture was originally derived from traditional fishing meth- ‘ods, such as the ‘acadias’ of Cate d'ivoire, West Africa (Welcomme, 1972), the ‘kathas’ of Bangladesh (Wahab and Kibria, 1994; Ahmed and Hambrey, 1999) and the ‘samarahs’ of ‘Cambodia (Shankar et al., 1998) (see also van Dam et al., 2002 for review). Dense masses of tree branches or bamboo are established in lakes, lagoons or rivers, and the fish are attracted by the provision of shelter from preda- tors, a suitable breeding habitat and the avail- ability of natural food, including periphyton (see Welcomme, Chapter 8 this volume for details) The potential of these unique fish-capturing tools for developing novel periphyton-based aquaculture systems was investigated by research groups in Africa and Asia Hem and Avit (1994) compared three 625 m? enclosures in Céte d'loire: one without substrate, one with a 100 m? acadja of a floating macrophyte, Echinochloa pyramidalis, and one with a 100 m? traditional acadja made of the usual tree branches. Fish migrated into these systems naturally through the 14-mm mesh sur- sounding nets. After 12 months, a total fish bio- mass of 11.7, 18.2 and 80.5 kg, respectively, wes harvested from the three enclosures. Tilapia, Sarotherodon melanotheron, con- tributed 80% of the total biomass in the enclo- sure with tree branches. Subsequent trials with different sizes of acadia (200-2500 m) yielded on average 1.8 tonnes/ha/year. Because of the high requirements for wood, additional trials8 ME. Azim etal. ‘were done with bamboo poles (10 sticks/m®, approximately 6 cm diameter), leading to aver- age vields of 8.3 tonnes/ha/year. Intensive cul- ture of two brackish water tilapia, Tilapia ‘guineensis and S. melanotheron, fed with pel- leted diet (30% protein) was also tried and it was reported that these two species were not suit- able for intensive culture but gave promising results in extensive acadja systems (Legendre etal., 1989) Although substrate-based aquaculture ‘was initiated in Africa, no pond-based trial was reported in the region. Some periphyton-based pond trials were carried outtin Asia in the1990s. Shrestha and Knud-Hansen (1994) conducted two experiments in Thailand where plastic sheets and bamboo poles (7.7 m? extra surface area per tank) were used vertically as substrates in conerete tanks (2.5 x 2 x 1.1 m3). Sex- reversed all-male Nile tilapia (Oreochromis niloticus) were stocked at a density of three fish per m? and grown over a 56-day period. Although there was evidence that the fish uti- lized periphyton from the substrates, there was no difference in net fish yield between tanks with and without plastic sheets as substrate, In another experiment, bamboo substrates resulted in greater net fish vield (3.43 g/m?/day) than plastic sheets (2.51 g/m®/day), but the contribution of bamboo substrate was not quan- tified due to absence of a treatment without bamboo substrate. The pioneering work on the Indian subcontinent was carried out at the Northwest Fisheries Extension Project (NFEP), Parbatipur aquaculture complex in Bangladesh by Farukcublslam (1996), who investigated the effect of introducing 40 vertical split bamboo panels (locally called chatai, each side 0.56 m2) on tilapia production in 80 m? trial ponds. Tilapia production in a 4-month culture period in ponds with and without bamboo chatai was 640 and 600 kg/ha, respectively. This differ- ence was not statistically significant. The bam- boo chatai was severely infested with snails, which competed with fish for food. In conclu- sion, these two initial experiments in Asia were not encouraging because of modest increases in fish production with additional substrate. ‘The second fish production trial conducted at the NFEP complex was more successful (NFEP, 1997). The trial was carried out in eight fertilized earthen ponds of 103-140 m? for a petiod of 168 days. Four ponds were provided ‘with bamboo trimmings inserted vertically into the pond bottom (17 poles/m?, diameter not mentioned) and four without substrates acted as controls. Juveniles of rohu, Labeo rohita (indi- vidual weight 11.5 g), were stocked at a density of one fish perm? in each pond. Net production in substrate and control ponds was 570 and 180 kg/ha, respectively; a threefold increase in production in the substrate treatment However, overall production of this trial was low because the trial commenced in August and continued till January. During the winter sea- son, low temperature (16°C) and short pho- toperiods limited fish production. Shankar et al. (1998) from India reported that the growth of Oreochromis mossambicus and Cyprinus carpio increased 48 and 50%, respectively, in fertized 1 m? cement tanks pro- vided with sugarcane bagasse (waste sugarcane fibre) as substrate compared to ponds without substrate during a 91-day culture period. Based on this result, a further investigation was carried out in three 25 m? cement cisterns (Ramesh etal., 1999), Sugarcane bagasse was suspended in cisterns fertilized with cow dung and urea, and only fertilized cisterns were used as control. Ata stocking density of 10,000 fish/ha, growth of C. carpio and L. rohita washigher by 47 and 48% respectively, compared to the conirol. The com- bined fish production was 1235 kg/ha in sub- strate ponds during a 133-day culture period. In recent years, the concepts of periphyton- based aquaculture were further tested and applied in monoculture (Keshavanath and Gangadhar, Chapter 13 this volume), polyculture (Azim and Wahab, Chapter 12 this volume), cage aquaculture (Huchette and Beveridge, Chapter 14 this volume}, shrimp culture (Tidwell and Bratvold, Chapter 15 this volume) and abalone culture (Kawamura et al., Chapter 16 this vol ume). 1.7. Outline/Structure of the Book To date, there has been no single monograph. ‘on periphyton, except for a chapter or a section in several books of the classic series ‘Limonology’ (e.g. Hutchinson, 1957; Wetzel, 1983a; Home and Goldman, 1994). In fact,Periphyton and Aquatic Production 9 ‘periphyton’ is widely referred to in the field of limnology but rarely considered in the aqua- culture literature. This book is the first mono- graph exclusively on periphyton, addressing the nature and function of periphyton in natu- raland aquatic farming systems and its possible role in enhancing aquatic food production. The book contributes to an increased understand- ing of the food web associated with periphyton and presents basic knowledge on using the periphyton loop for maximization of nutrient utilization in aquaculture systems. Exploring the suitability of periphyton in aquaculture ponds as natural fish food is a key objective; a considerable amount of material on periphyton ecology, its exploitation and management in open waters and other engineering systems has been assembled. Asa result, the book will be of interest to a wide audience, including stu- dents, teachers, researchers, extension work- ers and practitioners in the field of basic and applied aquatic ecology with a focus on limnol- ogy and aquaculture The chapters are contributed by renowned experts from around the world in their relevant fields. The chapters have been set out in a logi- cal sequence, The book begins with definitions of and synonyms for periphyton, followed by a brief summary of the role of periphyton in aquatic productivity in general, and in aquacul- ture ponds in particular (this chapter). The final chapter summarizes the state of the art and knowledge gaps with regard to periphyton research and discusses the potential of periphy- ton-based technology, with a particular focus on the emerging discipline of ecological aqua- culture (Costa-Pierce, 2002; Verdegem et al., Chapter 18 this volume). The remaining 16 chapters are organized around three main parts: Part I: Ecology (Chapters 2-5), Part Il: Exploitation (Chapters 6-9) and Part Ill: Management (Chapters 10-17) Part! (Ecology) provides an overview of the basic biology and ecology of periphyton com- ‘munities especially in freshwater ecosystems. Periphyton is categorized according to its mor- phological structure, biochemical composition and taxonomic diversity. Periphyton communi- ties in different habitats, together with coloniza~ tion and detachment processes, are reviewed in Chapter 2. Biomass and productivity of peri phyton communities vary greatly over short di tances and time spans, because constituent taxa can respond rapidly to changes in biotic and abiotic factors through colonization from the surrounding water and through growth. These issues are detailed in Chapter 3. Chapter 4 gives an in-depth insight into the role of peri- phyton in aquatic food webs. Energy and organic matter fluxes in general, growth regula- tion and recycling, community metabolism and links between periphyton and higher trophic levels are discussed here. While the previous chapters cover periphyton ecology in general, Chapter 5 particularly focuses on periphyton ecology in lakes and wetlands. The primary biotic and abiotic factors affecting lentic peri- phyton productivity are discussed. Part II (Exploitation) deals with the utiliza- tion of periphyton by aquatic animals, the func- tional morphology of periphyton grazers (fish) and exploitation of periphyton through fish- eries management and pollution control. Chapter 6 promotes an understanding of per- phyton systems from a systems ecology per- spective and analyses nutrient flows in periphyton-based pond culture. In this chapter, an attempt is made to synthesize the available knowledge on the processes underlying peri- phyton-based fish production in ponds into a quantitative framework that can help to under- stand the impact of periphyton substrates on the main nutrient cycles in ponds. Whether fish. are capable of grazing on periphyton depends mainly on their functional morphology as well as specialized adaptations for harvesting peri- phyton, Feeding ecology of periphyton grazers is discussed in Chapter 7. Brush park fisheries are fish-attracting devices (FAD) in open waters used in the tropics. Dense masses of tree branches or bamboo are established in lakes, lagoons or rivers and the fish are attracted by the provision of shelter from predators, a suitable breeding habitat and the availability of natural food. The form, function and per- formance, management strategies, socio-eco- nomic and environmental impacts of brush park fisheries are described in Chapter 8. Periphyton communities are regarded as efft- ient nutrient sinks and biological indicators in aguatic ecosystems. Chapter 9 focuses on the pros and cons of using periphyton as monitor- ing tools in natural and managed aquatic ecosystems. The structural and physiological10 MeE.Azim etal, descriptors of periphyton are critically exam- ined to evaluate their use as bioindicators. Part Ill (Management) reviews the applica: tion of periphyton in aquaculture systems, rang- ing from extensive pond culture to intensive recirculation systems, and from freshwater fin- fish to brackish water shellfish culture systems. Chapter 10 focuses on the interactions among periphyton, phytoplankton, water quality and different cultured fish species and recommends management strategies for periphyton-based earthen fishponds. Multivariate analysis statisti- cal tools were applied to understand the com- plex ecological processes in periphyton-based systems in this chapter. Heterotrophic microor- genisms dominate in biofilms in recirculation aquaculture systems. The similarities between ‘bacterial and periphytic biofilms are reviewed in Chapier 11. Attention is given to the use of periphyton to manage water quality, including the control of oxygen, ammonia and particulate and dissolved organic matter in culture systems. Pond polyculture is an age-old practice in Asia. The benefits of traditional polyculture can also be enjoyed in periphyton-based aquacul- ture systems. The provision of substrates may enhance the synergism among fish species. Periphyton-based Indian major carp polycul- ture and carp-freshwater prawn polyculture systems were developed in South and Southeast Asia and are described in Chapter 12. Ina thematic chapter (Chapter 13), per phyton-based aquaculture systems in India are described, based principally on research con- ucted in Mangalore, India. Besides tests of dit- ferent substrate materials with several indigenous fish species, this chapter covers periphyton biomass and digestibility and the effects of periphyton on fish health. Although aquaculture has been practised for several thousand years, cage aquaculture seems to be a relatively recent phenomenon. Today, an estimated 10% of annual freshwater finfish and 90% of marine finfish aquaculture production is derived from cages. ‘Biofouling’ is a common phenomenon in cage aquaculture, the term referring to the community of plants and animals that adheres to the frame and bag of floating cages (i.e. synonymous with ‘peri- phyton’). Caged herbivorous and omnivorous fish species can graze on net biofouling, thereby contributing fo growth and production. Chapter 14 presents the recent development of cage aquaculture with special attention on the exploitation of biofouling. Use of substrates in shellfish aquaculture has become popular in recent years. In Chapier 15, the utility of substrates in both marine and freshwater shrimp culture is reviewed. Freshwater shrimp exhibit strong temitoriality, which can be substantially mini- mized by providing layers of horizontal surface that increase the available territory per unit vol- ume of water. In contrast, marine shrimp do not exhibit such strong territorial behaviour. Therefore, surface additions in marine shimp systems have more commonly been vertical, to roughly mimic sea grass structures and enhance natural production associated with periphyton. Substrate orientation and densities, in relation to shrimp stocking densities are widely discussed. Marine benthic diatoms (peri- phyton) playa key role in the growth and devel- ‘opment of abalone. Substrates are the essential factor for larval settlement of abalone. Chapter 16 reviews abalone culture on both natural habitats and artificial surfaces. It also suggests the future use of periphyton in abalone hatch cries, describing various options for translating current knowledge into effective, practical techniques. Livelihood impacts and_ sustainability issues must be considered during the dissem- ination of any new aquaculture technol- ogy to farmers. The Aquaculture Research Programme (ARP) of the Department for International Development (DFID) in collabora- tion with the CARE Bangladesh-LIFE team ini- tiated tests of the periphyton technology in their development project and investigated the practical livelihoods impacts of this promising technology. Semi-structured interviews and participatory techniques, i.e. resource map- ping, activity calendars and ranking exercises, ‘were used to elicit the motivations, risks and livelihood outcomes from community members in two districts in Bangladesh. Chapter 17 pres- ents the results of the case study.Periphyton and Aquatic Production " References Ahmed, K.K, and Hambrey, J.B. (1999) Brush shelter: a recently introduced fishing method in the Kaptai reservoir fisheries in Bangladesh. ICLARM Quarterly NAGA 22(4), 20-23. ‘Azim, ME, (2001) The potential of periphyton-based aquaculture production systems. PhD thesis, Fish Culture ‘and Fisheries Group, Wageningen Institute of Animal Sciences, Wageningen University, The Netherlands, 219 pp. Aci, M.E., Wahab, M.A., van Dam, A.A, Beveridge, M.C.M., Mistein, A. and Verdegem, M.C.J. 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(1998) Constraints tothe use of plant fodder as fish feed in tropical small-cale tilapia culture SEioRe 20 overview. In: Kaushik, S.J. and Luquet, P. (eds) Fish Nutrition in Practice Inatine ‘National dela Recherche Agronomique, Les Colloques, No. 61, Paris, pp. 681-689.2 Periphyton Structure, Diversity and Colonization M.E. Azim andT. Asaeda Graduate School of Science and Engineering, Saitama University, Japan. 2.1. Introduction As with phytoplankton, periphyton can be found in almost every type of water body from small ponds to large oceans and in trophic con- Gitions that range from the most oligotrophic to the most eutrophic. Periphyton, in terms of structure, diversity, composition and the process of colonization, has received less attention from limnologists than phytoplankton. Since infor- mation is limited, the topic is reviewed here in general, regardless of habitats, with special attention to periphyton in ponds. Whenever the term periphyton or biofilm is used the defi- nitions from Chapter 1 apply. 2.2. Morphological Structure In essence, periphyton is a modified form of plankton, An individual alga, for example, might be found free swimming in the water column or attached to a substrate, or both, depending on. ‘when the observation is made and on the habitat. ‘The morphology of periphyton is therefore simi- lar to that of plankton, but with some additional adapiations which help periphyton to attach toa substrate and cope with the periphytic environ- ment. Algae show a wide range of morphological structures for adapting to a broad diversity of habitatsand substrates. Morphological character- istics differ among groups of algae and therefore different taxa show a variety of mechanisms for adapting to a sedentary life attached to sub- strates. Some of the more obvious morphologi- cal characteristics of periphytic organisms include stalks with sticky ends, sticky capsules, cushions or filaments, muscular suction pads, glue or simply clinging to the substrate. Attachment to sediment can also be achieved by rooting rhizoids and with a muscular foot (Reid and Wood, 1976). The size of periphytic algal organisms ranges from a single cell of about one thou- sandth of a millimeter in diameter toa large alga (a seaweed) up to 60 m in length. The shape varies from simple, non-motile single cells to motile, multicellular, filamentous structures. Non-motile forms of unicellular, colonial and fil ‘amentous algae attach to substrates by specially adapted cells and mucilaginous secretions, ‘Some taxa (e.g. Stigeoclonium) have morpho- logically distinct basal and filamentous cells, The basal cells form broad horizontal expanses of cells across the substratum surface and fila- ments develop vertically from them, Mucila ‘ous secretions can be amorphous for unicellular cyanobacteria and green algae, or organized into special pads, stalks or tubes for diatoms (Hoagland et al., 1982). Most of the unicellular and colonial cyano- phytes are non-motile whereas filamentous cyanophytes comprise both motile and non- motile forms. Gliding motility is by means of ‘©CAB International 2005. Periphyton: Ecology, Exploitation and Management (eds ME. Azim et al.) 5