ZOOLOGY Principles of Ecology Ecosystem Processes-II
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Paper No. : 12 Principles of Ecology Module : 33 Ecosystem Processes-II; Part-I Development Team Principal Investigator: Prof. Neeta Sehgal Head, Department of Zoology, University of Delhi Co-Principal Investigator: Prof. D.K. Singh Department of Zoology, University of Delhi Paper Coordinator: Prof. D.K. Singh Department of Zoology, University of Delhi Content Writer: Dr. Kapinder and Dr. Haren Ram Chiary Kirori Mal College, University of Delhi Content Reviewer: Prof. K.S. Rao Department of Botany, University of Delhi 1 Principles of Ecology ZOOLOGY Ecosystem Processes-II; Part-I Description of Module Subject Name ZOOLOGY Paper Name Zool 12 Principles of Ecology Module Name/Title Ecosystem Module Id M33 Ecosystem Processes-II; Part-I Keywords Ecological succession, mechanism of succession, theories of succession, climax characteristics Contents: 1. Learning Outcomes 2. Introduction 3. Causes of succession 4. Trends in succession 5. Types of Succession 5.1 Autotrophic succession 5.2 Heterotrophic succession 5.3 Primary succession 5.4 Secondary succession 5.5 Autogenic succession 5.6 Allogenic succession 5.7 Progressive succession 5.8 Retrogressive succession 6. Causes of Succession 7. Mechanism of Succession 8. Climax characteristics 9. Theories of Climax 9.1 Monoclimax theory 9.2 Polyclimax theory 9.3 Climax Pattern hypothesis 9.4 Information Theory 10. Summary 2 Principles of Ecology ZOOLOGY Ecosystem Processes-II; Part-I 1. Learning Outcomes After studying this module, you shall be able to: Understand the term Ecosystem. Learn the concept Ecological succession. Know different types of succession. Understand the causes and mechanism of succession. Learn the concept and theories of climax. 2. Introduction Succession is the ointment of time to the wounds of nature. F. Clement Organisms and the environment are interrelated and interact with each other so that flow of energy clearly defines the trophic structure, biotic diversity and material cycle within the system. The Term “Ecosystem” was coined by A.G. Tensley in 1935 who defined ecosystem as integration of living organisms in given area with the physical factors of the environment. It results in the flow of energy which causes development of distinctive trophic structure in the given system. An ecosystem is not a static system but it is dynamic in nature. Its structure and function changes with time in a sequential and orderly manner that can be predicted. This gradual and orderly change in the composition of species and processes of communities over time is called as ecological succession. The requirement of time for the process of succession differs with the nature of the climate, the structure and quality of soil. The term succession is usually applied for the directional and continuous change in composition and abundance of species following disturbance at a site or creation of a bare area. The process of succession generally results in a sequence of a number of communities which finally develops into a relatively stable and steady-state community. The directional changes of the communities results in biomass increase and change in the character and life form of species. 3 Principles of Ecology ZOOLOGY Ecosystem Processes-II; Part-I The whole sequence of communities in which one replaces the other in a given area is called sere. The first species established in the area form the pioneer stage exhibiting higher growth rate, short life span, smaller size and over reproduction. The final mature and stabilized stage of community is called climax which persists in the area until it is disturbed by some major natural or anthropogenic disturbance. Odum in 1963 defines ecological succession by explaining three parameters: 1. It is an orderly process in the community development which is directional and its result is predictable. 2. The process of ecological succession takes place due to change in environmental conditions caused by community itself. 3. The succession finally enters into steady state where maximum biomass and interaction of different organisms are continued per unit of energy flow in the system. 3. Causes of succession There are three main primary causes of succession: 1. Initial or Initiating causes. The initial cause of succession can be abiotic or biotic in nature. The abiotic factors include soil erosion and deposits, wind, temperature and fire. The biotic causes include activities of various living organisms. All these biotic or abiotic causes either produce bare areas or demolish the existing organisms in an area. 2. Ecesis or Continuing causes. Ecesis or continuing causes includes migration, aggregation, competition, reaction, etc. It results in successive waves of different populations due to various changes, primarily due to edaphic (soil) features of the area. 3. Stabilizing causes. These causes include several factors like climate of the area which result in the stabilization of the community. 4. Trends of succession Various trends which occur during the process of succession from initial (pioneer) to final (climax) stages include the following features: 4 Principles of Ecology ZOOLOGY Ecosystem Processes-II; Part-I 1. There is gradual change in the form of plants and animals in the area where succession is going on. It leads to increase in the species diversity. As the succession proceeds, the community structure and function become more complex. 2. As the succession continues, there is gradual increase in the living biomass and accumulation of dead organic matter. 3. The amount of chlorophyll continuously increases during the early stages of primary succession. Initially green pigment (chlorophyll) dominates but later diversity of other pigment also increases (carotenoid, xanthophylls etc). 4. In the early seral stage, production/respiration (P/R) ratio found more than 1 (production is more than respiration) but later in the climax stage, P/R ratio reaches to 1 (P/R = 1, production is equal to respiration). 5. In the beginning of succession, nutrients are allocated mostly in the soil, but in the later stage, maximum nutrients get allocated in the biomass and small amount present in the soil. As a result, biogeochemical cycle becomes more closed and effective. Food chain and food web also became more complex in the later stages of succession. 6. As the interrelation between different organisms became more complex, the role of detritivores becomes more important to recycle the material in the system. 7. The habitat for the community gets progressively modified into mesic condition from either dry or aquatic condition in the beginning of the succession. 8. The life cycle in the early seral stage was small and simple but in the later seral stages, it became more complex and longer. 9. The importance of macroenvironment becomes low as the succession proceeds toward climax stage. 10. Relationship among different species becomes mutalistic, however, other interactions such as competition, predation and amensalism occur to either regulate the population or prevent the invasion of outside elements. 11. In the early stage, dispersal of seeds and propagules occurs by wind, whereas, in the later stage it is mostly by animals. 5. Types of succession 5 Principles of Ecology ZOOLOGY Ecosystem Processes-II; Part-I Succession process is self propelled because each seral stage change the environment of the area in which it cannot renew itself. According to Margalef (1963) with progressive changes the succession attains maturity. However, Odum (1971) explained it as ecosystem strategy according to which there is a tendency of homeostasis in any ecosystem during unfavorable environment. On the basis of energy and nutrition relation, succession can be categorized into two types: 5.1: Autotrophic succession: It is the type of succession in which autotrophs (producers) dominate heterotrophs (consumers). This type of succession occurs in the area where the medium is rich in inorganic substances, thus the area is dominated by plants. As the producers are more in number, the rate of production becomes more the than rate of respiration. In the beginning, producers are present in majority but later the biomass of organism increase, as a result, the ratio of production and respiration becomes one. The diversity of organisms increases as the content of organic matter increases in the climax community. 5.2: Heterotrophic succession: In the early stages of heterotrophic succession, various heterotrophic organisms such as bacteria, fungi and actinomycetes are present in larger number. This type of succession starts in organic rich medium such as the river or streams which are highly polluted with organic matter (sewage), or in the small pools having large quantities of leaf litter. As the heterotrophic succession is initially dominated by heterotrophs, so the rate of respiration becomes greater than production. It can be well studied in the heterotrophic communities of the organisms which are involved in the decomposition processes such as dead parts of the plant, animal remains and wastes etc. In these communities, plants and animals succeed each other in colonization and replacement process. In this type of succession, fungi and some invertebrates dominate the area and they feed on dead and decaying organic matter. High amount of energy and nutrients are available in the beginning of the succession and gradually decline as succession proceeds. In addition to the autotrophic and heterotrophic succession, following types of succession is also present: 5.3: Primary succession: