Environmental Biology I: Ecology as an Interdisciplinary Science, the Origin of Life and Speciation, Human Ecology, Ecosystem Structure and Function, Energy Flow, Succession, Ecotones, Niches, Stability and Services, the Types of Ecosystem and the Biomes
2. Ecosystem structure and function: energy flow, food chains and webs, pyramids and cycles
An ecosystem, a term introduced by Tansley in 1935, has abiotic components (energy, climate, water, inorganic substances and organic compounds such as humus) and biotic components: producers (autotrophs), consumers (herbivores, carnivores, omnivores) and decomposers (bacteria and fungi, with detritivores). Its two basic functions are the one-way flow of energy and the cycling of matter. Gross primary productivity (GPP) is the total rate of photosynthesis; net primary productivity (NPP) is what remains after the producers' own respiration, NPP = GPP − R, and it is the energy actually available to consumers. Tropical rainforests, estuaries, coral reefs and swamps have the highest NPP per unit area, and the open ocean and deserts the lowest, though the ocean's great area makes it a large total producer.
Odum's energy flow models show the same principle at increasing detail. The single-channel model traces energy from producers through herbivores to carnivores, with losses to respiration at every step. The Y-shaped model splits the flow into a grazing food chain (plants to herbivores to carnivores) and a detritus food chain (dead organic matter to decomposers and detritivores to their predators); in most forests and many aquatic systems far more energy passes through the detritus chain. The universal model applies to any living component, showing ingested energy divided into assimilation and egestion, and assimilation into respiration and production. Food chains interlock into food webs, which give consumers alternative prey and so stabilise the community. Because only about 10% of energy passes from one level to the next (Lindeman, 1942), if producers fix 20,000 kJ, herbivores receive about 2,000 kJ and carnivores about 200 kJ, which is why food chains rarely exceed four or five links.
| Ecological pyramid | Always upright? | Inverted example |
|---|---|---|
| Numbers | No | A single large tree supporting thousands of insects; a host with many parasites |
| Biomass (standing crop) | No | Open ocean, where a small standing crop of fast-dividing phytoplankton supports a larger biomass of zooplankton |
| Energy (productivity) | Yes | None, because energy is lost at every transfer |
Matter, unlike energy, cycles. Biogeochemical cycles are of two broad types: gaseous cycles, such as those of carbon, nitrogen and oxygen, have their main reservoir in the atmosphere or ocean and are relatively well buffered; sedimentary cycles, such as that of phosphorus, have their reservoir in rocks and sediments, turn over slowly and are easily disrupted, which is why phosphorus lost to the sea is effectively lost to the land for geological time. Each cycle has a large, slow reservoir pool and a smaller, active exchange pool between organisms and their surroundings.
3. Ecological succession
Succession is the orderly, directional change in the species composition of a community over time. Primary succession begins on a surface never before colonised, such as bare rock, a lava flow or a new sand dune, and is slow because soil must first form; secondary succession follows disturbance of an existing community, such as an abandoned field or a burnt forest, where soil remains, and is faster. On bare rock (a lithosere, a kind of xerosere) the sequence runs from crustose lichens, which begin weathering, to foliose lichens, mosses, herbs, shrubs and finally forest; in water (a hydrosere) from phytoplankton to submerged plants, floating plants, reed swamp, sedge meadow, woodland and forest, as sediment and organic matter fill the basin. Changes caused by the organisms themselves are autogenic; those caused by outside forces such as siltation or climate are allogenic.
The end point is the climax community, in equilibrium with its environment. Clements's monoclimax theory held that each climatic region has a single climax; Tansley's polyclimax theory allowed several climaxes set by soil, topography and fire; and Whittaker's climax-pattern hypothesis sees a continuous pattern of climax communities tracking environmental gradients. Connell and Slatyer (1977) proposed three mechanisms by which early species affect later ones: facilitation, tolerance and inhibition. Odum (1969) summarised the trends of a developing ecosystem: biomass and organic matter increase, species diversity and structural complexity increase, food chains become web-like and detritus-based, nutrient cycles become more closed, K-selected species replace r-selected ones, and the ratio of gross production to community respiration (P/R) moves towards one.
4. Ecotones, edge effects, habitats and niches, ecosystem stability and ecosystem services
An ecotone is the transition zone between two communities, such as a forest edge meeting grassland, or a mangrove between land and sea. It often holds species of both communities plus some found only there, and the resulting tendency for greater diversity and density at the junction is the edge effect; species that prefer such zones are edge species. Fragmentation multiplies edges, which benefits edge species but harms interior species that need large blocks of undisturbed habitat. The habitat is where an organism lives, its address; the niche is its role, its profession. Grinnell (1917) used niche mainly in the spatial sense, Elton (1927) for an organism's functional place in the community, and Hutchinson (1957) defined it as an n-dimensional hypervolume of all the conditions and resources in which a species can persist. The fundamental niche is that hypervolume without competitors; the realised niche is the narrower part actually occupied in their presence. By Gause's competitive exclusion principle, two species with identical niches cannot coexist indefinitely.
Stability has two components: resistance, the ability to withstand a disturbance without changing, and resilience, the ability to return to the original state after one. The diversity-stability hypothesis of MacArthur (1955) and Elton (1958) holds that more diverse communities, with more pathways for energy, are more stable. Robert May (1973) showed mathematically that randomly assembled complex systems tend to be less stable, and the modern view is that real communities are stabilised not by complexity as such but by particular structures, such as many weak interactions, functional redundancy among species and the insurance that different species respond differently to change. Other factors affecting stability are the presence of keystone species, the frequency and intensity of disturbance, and the connectivity of habitat.
| Ecosystem service (Millennium Ecosystem Assessment, 2005) | Examples |
|---|---|
| Provisioning | Food, fresh water, timber, fibre, fuel, genetic resources, medicines |
| Regulating | Climate regulation, flood control, water purification, pollination, pest and disease control |
| Cultural | Recreation, tourism, spiritual and religious values, aesthetic inspiration, education |
| Supporting | Soil formation, nutrient cycling, primary production, which underpin all the others |
Costanza and colleagues (1997) estimated the value of the world's ecosystem services at about 33 trillion US dollars a year, more than the global economic output of the time, a figure intended less as a price than as a demonstration that services treated as free are not.
5. The basis of classification and the types of ecosystem
Ecosystems are classified by habitat (terrestrial or aquatic, and within aquatic, freshwater, estuarine or marine), by origin (natural or human-made, such as croplands, plantations and reservoirs), by the dominant vegetation or physiognomy, and, in Odum's energetic scheme, by the source of energy: unsubsidised natural solar-powered systems such as the open ocean, naturally subsidised ones such as estuaries enriched by tides, human-subsidised solar-powered ones such as farms, and fuel-powered urban-industrial systems.
| Ecosystem type | Defining features | Indian example |
|---|---|---|
| Hot desert | Rainfall usually below 250 mm, high temperature range; plants with succulence, CAM photosynthesis and deep roots | Thar Desert, Rajasthan |
| Cold desert | High altitude, very low temperature and precipitation, strong radiation | Ladakh, Lahaul-Spiti |
| Forest | Tree-dominated; stratified canopy; the largest store of terrestrial carbon | Tropical evergreen forests of the Western Ghats; sal forests of central India |
| Rangeland | Grasslands, shrublands and savannas used for grazing; productivity limited by rainfall and grazing pressure | Banni grassland, Kachchh; shola grasslands of the Nilgiris |
| Wetland | Marsh, fen, peatland or water, natural or artificial, permanent or temporary, fresh, brackish or salt, including marine water not deeper than six metres at low tide (Ramsar definition) | Chilika, Keoladeo, Loktak, the East Kolkata Wetlands |
| Lotic (running water) | Current, high oxygen, organisms with attachment and streamlining; changes from source to mouth (river continuum concept, Vannote and colleagues, 1980) | Himalayan torrents to the lower Ganga |
| Lentic (standing water) | Littoral, limnetic, profundal and benthic zones; temperate lakes stratify into epilimnion, thermocline and hypolimnion, overturning in spring and autumn | Dal Lake, Wular Lake, village ponds |
| Estuarine (mangrove) | Mixing of fresh and sea water, fluctuating salinity, very high productivity; mangroves with pneumatophores and vivipary | The Sundarbans; Bhitarkanika |
| Oceanic | Neritic and oceanic zones; a sunlit photic zone over a dark aphotic zone; productivity concentrated in upwelling and coastal waters | The Arabian Sea and Bay of Bengal, with seasonal upwelling off the south-west coast |
6. Biomes: concept, classification and distribution
A biome is a large regional community of plants and animals with a characteristic physiognomy, determined above all by climate: Whittaker's diagram places the world's biomes on axes of mean annual temperature and precipitation. Biomes grade into each other along latitude, and a similar zonation repeats with altitude on high mountains. The syllabus spells chaparral as "Chapparal"; the correct spelling has one p.
| Biome | Climate | Vegetation, soil and fauna |
|---|---|---|
| Tundra | Very cold, short growing season of about two months, low precipitation; permafrost | Mosses, lichens, sedges, dwarf shrubs; no trees; caribou, arctic fox, migratory birds |
| Taiga (boreal forest) | Long cold winters, short summers, moderate precipitation | Evergreen conifers such as spruce, fir and pine; acidic podzols; moose, lynx, wolves |
| Temperate grassland | Hot summers, cold winters, 250 to 750 mm rain | Grasses on deep, fertile chernozems; the prairies, steppes, pampas and veld; bison and grazing herds |
| Temperate deciduous forest | Four distinct seasons, rain spread through the year | Broad-leaved trees such as oak, maple and beech that shed leaves in autumn; fertile brown forest soils |
| Highland icy alpine | Above the tree line on high mountains; cold, windy, intense ultraviolet | Cushion plants, alpine meadows, rhododendron scrub; snow leopard, ibex, yak in the Himalaya |
| Chaparral (Mediterranean scrub) | Hot, dry summers and mild, wet winters | Dense evergreen sclerophyllous shrubs adapted to fire; California, the Mediterranean basin, central Chile, the Cape of South Africa, south-west Australia |
| Savanna | Warm all year with distinct wet and dry seasons | Grassland with scattered trees, maintained by fire and grazing; the great herds and predators of East Africa |
| Tropical rainforest | Hot and wet all year, often over 2,000 mm of rain | Multi-layered evergreen forest with the highest species richness on land; most nutrients are held in the vegetation, and the leached soils are poor once it is cleared |
Key takeaways
- Life probably arose by chemical evolution (Oparin-Haldane, Miller-Urey); species form by allopatric, peripatric, parapatric and sympatric routes; cities are heterotrophic ecosystems.
- NPP = GPP − R; energy flows one way through grazing and detritus chains, about 10% passing each level, so the energy pyramid is always upright while those of number and biomass can invert.
- Succession runs from pioneers (crustose lichens on rock, phytoplankton in water) to a climax, with biomass, diversity and closed nutrient cycles increasing and P/R approaching one.
- Ecotones show the edge effect; Hutchinson's niche is an n-dimensional hypervolume, fundamental or realised; stability combines resistance and resilience; services are provisioning, regulating, cultural and supporting.
- Ecosystem types run from hot and cold deserts to lotic, lentic, estuarine and oceanic waters, and biomes from tundra and taiga to chaparral (the syllabus's "Chapparal"), savanna and tropical rainforest, each set chiefly by temperature and rainfall.
Practice questions (10)
Attempt each one before opening the answer. Every explanation names the tempting wrong option as well as the right one, because that is where marks are lost.
Which ecological pyramid is always upright?
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Answer: D — The pyramid of energy
Energy is lost as heat at every transfer, so each level must have less energy flow than the one below it. The pyramid of numbers inverts under a single large tree and that of biomass in the open ocean, where fast-turning phytoplankton support a larger standing crop of zooplankton.Producers in a grassland fix 20,000 kJ of energy. Assuming 10% transfer between successive trophic levels, how many kJ reach the secondary consumers?
Numerical answer — type the value.
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Answer: 200
Primary consumers (herbivores) receive 10% of 20,000 = 2,000 kJ, and secondary consumers (the carnivores that eat them) receive 10% of that, 200 kJ. Stopping at 2,000 kJ confuses primary with secondary consumers.Hutchinson's concept of the ecological niche is best described as
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Answer: A — an n-dimensional hypervolume of the conditions and resources within which a species can persist
Hutchinson (1957) defined the niche as a multidimensional space of all the environmental variables a species tolerates and the resources it uses. The physical place is the habitat, the transition zone is an ecotone, and a purely trophic position is closer to Elton's functional niche.Which of the following are trends expected as an ecosystem develops towards climax, according to Odum? Select all that apply.
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Answer: A — Total biomass and organic matter increase; C — The ratio of gross production to community respiration approaches one
In a maturing ecosystem biomass accumulates and production comes into balance with respiration, so P/R moves towards one. Nutrient cycles become more closed, conserving nutrients, and K-selected species replace the r-selected pioneers.A biome with hot, dry summers, mild, wet winters and dense, fire-adapted evergreen sclerophyllous shrubs is the
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Answer: B — chaparral
That is the Mediterranean climate of the chaparral, found in California, the Mediterranean basin, central Chile, the Cape and south-west Australia; the syllabus spells it "Chapparal". Savanna has a warm wet-and-dry climate with grass and scattered trees, and taiga and tundra are cold biomes.Assertion (A): In the open ocean the pyramid of biomass can be inverted. Reason (R): Phytoplankton reproduce so fast that a small standing crop can support a larger biomass of zooplankton. Choose the correct option.
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Answer: A — Both A and R are true, and R is the correct explanation of A
Both are true and R explains A. Biomass is a snapshot of standing crop, not a rate; phytoplankton turn over in days, so their small standing stock produces enough to feed a larger, slower-turning mass of consumers. The pyramid of energy, which measures rates, stays upright.In the Millennium Ecosystem Assessment, soil formation, nutrient cycling and primary production are classed as
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Answer: B — supporting services
Supporting services are the underlying processes on which all others depend. Provisioning services are products such as food and timber, regulating services include climate regulation and pollination, and cultural services are non-material benefits such as recreation and spiritual value.Which statements about thermal stratification in a temperate lake are correct? Select all that apply.
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Answer: A — The hypolimnion may become depleted of oxygen during summer stratification; B — The thermocline is the layer in which temperature falls rapidly with depth; D — Spring and autumn overturns mix oxygen and nutrients through the water column
In summer a warm, light epilimnion floats over a cold, dense hypolimnion, separated by the thermocline, and decomposition can strip the isolated hypolimnion of oxygen. When surface water cools or warms to about 4 °C in autumn and spring, density differences vanish and the lake mixes. The epilimnion is the warm upper layer, not the bottom one.Speciation that occurs when a population is split by a geographical barrier such as a mountain range is
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Answer: A — allopatric speciation
Allopatric means "other homeland": the barrier stops gene flow and the isolated populations diverge. Sympatric speciation happens within one area, often through polyploidy, and parapatric speciation happens across a gradient between adjacent populations still in contact.A forest has a gross primary productivity of 25,000 kcal/m²/yr, and its plants use 60% of this in respiration. If primary consumers obtain 10% of the net primary productivity, how many kcal/m²/yr do they obtain?
Numerical answer — type the value.
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Answer: 1000
NPP = GPP − R = 25,000 − 0.6 × 25,000 = 10,000 kcal/m²/yr, and 10% of that is 1,000 kcal/m²/yr. Taking 10% of the GPP (2,500) ignores the plants' own respiration, which is exactly the part consumers can never obtain.