A tabular model for ecosystem development

A more general and complete summary of important changes in community structure and function in the sere, as revealed by the study of the large, open systems of nature, is showing this post....

Expected trends in the gradient from youth to maturity are grouped under several headings. Although ecologists have studied succession in many parts of the world most of the emphasis to date has been on the descriptive aspects such as the qualitative changes in species structure. Only recently have the functional aspects of succession also been considered. Consequently, some of the items listed in this post must be considered hypothetical in the sense that they are based on good experimental or theoretical evidence, but have not been verified by adequate data from the field. Five aspects seem most significant require a bit more explanation as follows.

The kinds of plants and animals that change continuously with succession. Those species that are important in the pioneer stages are not likely to be important in the climax. When the density of species in a sere is plotted against time, a characteristic stair – step graph is obtained, as illustrated in this post. Such a pattern usually is apparent whether we are considering a specific taxonomic group, such as birds, or a trophic group, such as herbivores or producers. Typically, some species in the gradient have wider tolerances or niche preferences than others and, therefore, persist over a longer period of time. Thus, in the terrestrial succession pictured in thispost pine trees and cardinals persist throush longer periods of time than do most of the other species. In general, the more species in the group (whether taxonomic or ecological) that are geographically available for colonization, the more restricted will be the occurrence of each species in the time sequence. This kind of regulatory adjustment is the result of competition coexistence interactions discussed in the next post.

Biomass and the standing crop of organic matter increase with succession. In both aquatic and terrestrial environments the total amount of living matter (biomass) and decomposing organic materials tend to increase with time. Also, many soluble substances accumulate; these include sugars, amino acids, and many organic products of microbial decomposition. These liquid products that leak out from the bodies of organisms are often collectively known as extra metabolites. Some of these substances provide food for microorganisms, and perhaps also for microorganisms. Other   substances are equally important in that they may act as inhibitors (antibiotics) or as growth promoters (as, for example, vitamins), sub stances produced by one organism may inhibit the further growth of that species (thus providing population self regulation, or they may act on completely different species. This was dramatically drought to our attention by the discovery of penicillin and other bacterial antibiotics produced by fungi. In other cases, increasing organic matter stimulates the growth of bacteria that manufacture vitamin B12, a necessary growth promoter for many animals (many are unable to manufacture this and other vitamins themselves). Where extra metabolites do prove to be regulatory, we would be justified in calling these substances environmental hormones since by definition a hormone is a ‘’chemical regulator.’’ Chemical regulation is one way of achieving community stability as the climax is approached, because the physical as well as the chemical perturbations (as, for example, light and water relations) are buffered by a large organic structure. There is no question that the increase in amount of and the change in organic structure are two of the main factors bringing about the change in species during ecological development.

The diversity of species tends to increase with succession. Initially this is the case, although it is not clear from the present data that  the change in variety of taxa follows the same pattern in all ecosystems. Increase in diversity of heterotrophs is especially striking; the variety of microorganisms and heterotrophic plants and animals is likely to be much greater in the later stages of succession than in the early stages. Maximum diversity of autotrophs in many ecosystems seems to be reached earlier in succession. The interplay of opposite trend makes it difficult to generalize in regard to diversity. The increase in size of individual organisms and the increase in competition tend to reduce diversity, while the increase in organic structure and variety of niches tends to increase it. As we have already pointed in the discussion of diversity in this post there may be an optimum level of diversity for a given energy flow pattern. We can state that, in general, rapid growth seral stages  will tend to have a low diversity on the order of 0.1 or 0.2 on the scale used in this post while mature stages will tend to have a higher level on the order of 0.7 or 0.8 unless there is a large energy subsidy that counteracts this patter.

A decrease in net community production and a corresponding increase in community respiration are two of the most striking and important trends in succession. These changes in community metabolisms are shown graphically in this post which compares ecosystem development in a small laboratory microcosm and in a large natural forest. Total production (PG ) increases faster than energy expenditure (P) at first, so a large net production (PN ) results in a rapid increase in biomass (B). gradually, equilibrium is established, in about 100 days in the microcosm and 100 or more years in the forest. Perhaps the best way to picture this overall trend is as follows; species, biomass, and the P/R ratio continue to change long after the maximum gross primary production possible for the site has been achieved. As one evidence for this we may cite the situation in regard to leaves in a terrestrial broadleaved succession. Agricultural scientists have repeatedly found that maximum productivity of broad leaved crops occurs when the leaf surface area exposed to the incoming light from above is about 4 or 5 times the surface area of the ground. Any increase in leaves beyond this level does not increase the photosynthetic rate per square mater, since increased shading cancels any advantage that might accrue from increased photosynthetic tissue. In fact, the increased respiration of the extra leaves that do not receive adequate light may reduce the net production of the crop. In a forest the leaf area apparently continues to increase far beyond that limit experimentally shown to increase gross production, since leaf area per ground is often 10 or more in an old forest. Since forests are among, the most successful of ecosystems with a long geological history of survival, we may well consider the possibility that the extra leaves have other important functions in the ecosystem in addition to production of food. The undoubtedly help moderature and moisture and provide reserves that are important during periods of climatic stress or insect of climatic stress or insect or disease attack.  

A tabular model for ecological succession of the autogenic, autotrophic type

Ecosystem characteristic
Trend in ecological development  early stage to climax
youth to maturity
growth stage to steady state

Community structure
Species composition
Changes rapidly at first, then more gradually
Size of individuals
Tends to increase
Number species of autotrophs
Increases in primary and often early in secondary succession; may decline in older stages as size of individuals increases
Number species of heterotrophs
Increases until relatively late in the sere
Species diversity
Increases initially, then becomes stabilized or declines in older stages as size of individual increases
Nonliving organic matter

Energy flow (community metabolism)
Gross production (P)
Increases during early phase of primary succession; little or no increase during secondary succession
Net community production (yield)
Community respiration (R)
P/R ratio
P>R to P = R
P/B ratio
B/P and B/R ratios (biomass supported/ unit energy)
Food chains
From linear chains to more complex food webs

Biogeochemical cycles
Mineral cycles
Become more closed
Turnover time
Role of detritus
Nutrient conservation

Natural selection and regulation

Growth form
From r – selection (rapid growth) to K – selection (feedback control)a
Quality of biotic components
Increasing specialization
Life cycles
Length and complexity increases
Symbiosis (living together)
Increasingly mutualistic
Overall stability

Evolution of the ecosystem

     As in the case with short term development, as described earlier in this post, the long term evolution of ecosystems is shaped by the interaction of allogenic geological and climatic changes and autogenic processes resulting from the activities of the living components of the ecosystem. In a broad sense the ‘’strategy’’ of long term evolutionary development is the same as that of short term ecological succession, namely, increased control of, or homeostasis with, the physical environment in the sense of achieving maximum protection from its perturbations.
     Although we may never know exactly how life began on earth, the generally accepted theory is that the first living things were tiny anaerobic (living without free oxygen) heterotrophs that lived on organic matter synthesized by abiotic processes. The first successional development then, may been more like the hay infusion culture model (see that post) than the autotrophic culture model. The atmo phere at the time of the origin of life 3 billion years ago containe nitrogen, hydrogen, carbon dioxide, water vapor, but little or oxygen. It also contained carbon monoxide, chlorine, and hydrogen sulfide in quantities that would be poisonous to much of present day life. The composition of the atmosphere in those early days was largely determined by the gaseous stuff that comes out of volcanos. The geologist would speak of this as ‘’atmospheric formation by crusts outgassing.’’ The earth’s early reducing atmosphere (a term to contral with oxygenic atmosphere) may have been similar to that now found on venus or jupiter. Because of the lack of gaseous oxygen there was no ozone laver, as there is now. Molecular oxygen o2 acted on b short – wave untraviolet radiation produces ozone, or 03 which in turshields out the deadly radiation. Thus, at first, life could existonly inshielded by water or other barrers, but straange to say it was the short waved radiation that is thought to have created a chemical evolution leading to complex organic molecules such as amino acide that became the building blocks of life. This synthesis also provided food for the first organisms.
      For millions of years life apparently remained as only a tiny foot hold, limited in habitat and energy source, in a violent physical world. The big change began with the appearance of the first photosynthetion algae which were able to make food from simple inorganic substances and which released gaseous oxygen as a by product. As the oxygen diffused into the atmosphere, the ozone shield developed and life could then spread to all parts of the globe, and there followed an almost explosive evolution of increasingly complex aerobic organisms. The broad pattern of the evolution of organisms of organisms and the oxygenic atmosphere that make the biosphere absolutely uniqe in our solar system is shown in this post. Over long stretches of time production exceeded respiration (P/R > 1) so oxygen increased and CO2 decreased. Our fossil fuels were also formed during periods when P exceeded R by a wide margin.
      Incidentally, I can think of no batter way to dramatize man’s dependence on his environment and to become a wise custodian of this frail earth than to recount how our atmosphere came into being, emphasizing, of course, that it was built by microorganisms, not by men. I think the story of our air should be told to every school child every citizen. It is a fascinating drama of living history with enough mystery and potential tragedy to intrigue teacher and pupil alike. It is a subject lends itself to student participation in learning since the possibilities for study projects, artwork, plays, and the like are unlimited. Berkner and marshall have writter both a population account (1966) and a more technical treatise (1964) that provide good reference.

The significance of ecological development to environmental management by man

     The  aciples  of ecological ecosystem development are of the greatest importance to mankind. Man must have early succcessional stages as a continuous source of food and other organic products, since he must have a large net primary production to harvest; in the climax community, because production is mostly consumed by respiration (plant and animal), net communitu production in an annual cycle may be zero. On the other hand, the stability of the climax and its ability to buffer and control physical forces (such as water temperature) are desirable characteristics from the viewpoint of the human population. The only way man can have both a productive and a stable environment is to consure that a good mixture of early and mature successional stages are maintained, with interchanges of energy and materials. Excess food produced in young communities helps feed older stages that in return supply regenerated nutrients and help buffer the extreme of weather (storms, floods and so on).

   In the most stable and productive natural situation there is usually such a combination of successional stages for example, in areas such as the inland sea of japan or long island sound, the uoung communities of plankton  feed lder, more stable communities on the rocks and on the bottom (benthic communities) the large biomass structure and diversity of the benthic communities provide not only habitat and shelter for life hisory stages of pelagic forms but also regenrtated nutrients necessary for continued productivity of the plankton. A similar, favorable situation exists in many terrestrial landscapes where productive croplands on the plains are intermingled with diverse forests and orchards on the hills and mountains. The  crop fields are, ecologically speaking, ‘’young nature’’ in that they are maintained as such by the constant labor of the farmer and his machines. The forests represent older, more diverse, and selfsustaining communities that have lower rates of net production but do not require the contant attention of man. It is important that both types of ecosystems be considered together in proper relation. If the forests are destroyed merely for the temporary gain in wood production, water and soul may wash down from the slopes and reduce the peoductivity of the plains. Ruins of civilizations and man made deserts in various parts of the world stand as evidence that man has not been fully aware of his need for protective as well as productive environments. Mature systems have other values to mankind in addition to products; they should not be considered as crops in the sense of wheat or corn. The conservationist speaks of a policy of balancing contradictory needs as ‘’multiple use,’’ but in the past he has found it difficult to translate long term values into monetary units. Consequently, too often the possibilities for immediate economic gain in harvest overrides what later turns out to be a more important value.

    To illustrate thesee difficulties, let us consider the controversy over national forest management policy that received considerable public attention in the early 1970s.for the most part national forests have been managed on a ‘’selective cut’’ basis; that is, selected trees, inclding a portion of mature trees that are no longer growing are removed periodically leaving the stand more or less intact to serve other uses (recreation, soil and water stabilization, and so on) and leaving room for younger trees to grow faster.  As the demand for paper and other wood products became acute threr was pressure for harvesting on a ‘’clear cut and replant’’ cycle, since the yield would then be greater and subsequent rate of net production incrrased. But right after the clear cut the system would be subject to various disorders such as soul erosion and nutrient loss; the cost of taking care of these problems could cancel out the value of extra wood yield. Thus, both plans have advantages and disadvantages. A sensible solution to the dilemma would be to vary the management according to the site capability. Where topography is steep and soil thin, or where the vegetation is botanically unique or of great scenic beauty, a selective cut plan would be best in the long run. Where topography is more level, the soil deep and stable, and the species caplable of repid regrowth, then a clear cut procedure could be a desirable choice.

In essence there are only two basic ways to meet the problem of youth and maturity in the landscape. One would be to maintain intermediate states as naturally occurs in pulse stabilized systems, and the other would be to compartmentalize or ‘’zome’’ the landcape so as to have separate areas primarily managed for production and for protedtion. Both requure that society adopts regional land use plans, an idea whose time is coming.

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