Jugami Biology

Biology · Lesson 5

Ecology and Ecosystems

Who eats whom, where the nutrients go, and why one missing sea star can change a whole shore.

Ecology is the study of how living things interact with each other and with their surroundings. It asks why a species lives where it does, how many of it there are, and what happens to everything else when that number changes.

You will see energy from Photosynthesis and Respiration flow through whole ecosystems, while carbon and nitrogen go round in cycles.

Ink sketch of an energy pyramid: producers at the base hold 10,000 units of energy, plant eaters 1,000, meat eaters 100 and top predators 10, with the sun shining on the base.10,000producers1,000plant eaters100meat eaters10top predatorsfig. 1: energy at each level

1. Levels of organization

Ecologists work at several scales, each one containing the one before:

  1. OrganismOne living thing, such as a single sea star.
  2. PopulationAll the members of one species in an area: every ochre sea star on a stretch of shore.
  3. CommunityAll the populations of different species living and interacting in the same place.
  4. EcosystemA community plus its nonliving surroundings: rock, water, light, temperature and nutrients.
  5. BiomeA large region with a similar climate and similar kinds of communities, such as tundra, desert or tropical rainforest.
  6. BiosphereEvery part of Earth where life exists, from the deep sea to the upper air.

2. Producers, consumers and decomposers

Producers (autotrophs) make their own food. Most use photosynthesis: plants on land, and algae and cyanobacteria in water. A few bacteria near deep-sea vents use chemical energy instead.

Consumers (heterotrophs) get energy by eating other organisms. Primary consumers eat producers, secondary consumers eat primary consumers, and so on. Each feeding step is a trophic level. Many animals feed at more than one level: a bear that eats berries and salmon is both a primary and a higher-level consumer.

Decomposers, mainly bacteria and fungi, break down dead matter and waste. Without them, nutrients would stay locked in dead bodies and the soil would run out.

3. Food chains and food webs

A food chain is a single path of who eats whom: seaweed → limpet → sea star. Real communities are tangled, because most species eat, and are eaten by, several others. A food web shows all of those links at once.

In 1963 the ecologist Robert Paine began a simple experiment on the rocky shore at Mukkaw Bay, Washington. He kept prying ochre sea stars off one stretch of rock and throwing them back into the sea, and left a nearby stretch alone. The plot without sea stars was soon covered by California mussels, which crowded out barnacles, seaweeds, chitons and limpets. Within a few years the number of species there had fallen from 15 to 8. Paine later called a species like this, whose effect is far larger than its numbers suggest, a keystone species.

Food web of a rocky shore in Washington State, after Paine (1966)Plankton feed acorn barnacles and California mussels. Seaweeds feed chitons and limpets. Whelks eat barnacles. The ochre sea star eats barnacles, mussels, chitons, limpets and whelks. Mussels and barnacles compete for space on the rock.Ochre sea starPisaster ochraceusWhelkNucella (Thais)Acorn barnaclesBalanusCalifornia musselMytilus californianusChitonsKatharinaLimpetsAcmaeaPlanktondrifting in the waterSeaweedsand other algaeArrows point from food to eater.Dashed: compete for rock space

What Paine found: with the sea star gone, mussels took over the rock. Barnacles, seaweeds, chitons and limpets lost space, and the plot fell from 15 species to 8.

Turn on JavaScript to remove other species and see what the web predicts.

Who eats whom

  • Plankton feeds barnacles and mussels.
  • Seaweeds feed chitons and limpets.
  • Whelks eat barnacles.
  • The sea star eats barnacles, mussels, chitons, limpets and whelks.
  • Mussels and barnacles compete for space on the rock.

This is a qualitative sketch: it shows the likely direction of change, not amounts. Only the sea star removal was tested in the field. The other outcomes are predictions read off the arrows.

Trophic cascades. Removing a top predator can ripple down through several levels. Gray wolves were wiped out of Yellowstone National Park by the 1920s and reintroduced in 1995 and 1996. Elk numbers fell in the years that followed, and some studies report that willows and aspens began to recover where elk browsing eased. Other researchers argue the effect is weaker or patchier than the popular story suggests, because drought, hunting, bears and cougars also affect elk, and aspen recovery varies from place to place. Claims that wolves changed the course of Yellowstone's rivers are especially debated. Wolves are back in the park's food web, but ecologists are still sorting out which of the changes they caused.

4. Energy pyramids and the 10% rule

Energy enters most ecosystems as sunlight captured by producers and leaves as heat. Only a fraction of the energy at one trophic level ends up in the bodies of the next. The rest is used for the organisms' own respiration and lost as heat, or it is in parts that are never eaten or digested.

On average about 10% passes from one level to the next, which is why the hero sketch at the top of this page drops from 10,000 to 1,000 to 100 to 10. This is a rough average, not a law: measured transfer efficiencies vary, roughly between 5% and 20%, depending on the ecosystem and the animals involved.

The rule explains why food chains rarely have more than four or five levels and why top predators are scarce. Pyramids of numbers or biomass can sometimes be upside down (one oak tree can feed thousands of caterpillars), but a pyramid of energy never is.

5. Carbon and nitrogen cycles

Energy flows through an ecosystem in one direction, but matter is recycled. Click a pink step in either diagram, or use the buttons, to read about it.

The carbon cycle

The carbon cycleCarbon dioxide moves from the atmosphere into plants and algae by photosynthesis. Eating passes carbon to animals. Respiration by plants, animals and decomposers returns CO₂ to the air. Dead matter goes to decomposers, and some is buried over millions of years to form fossil fuels; burning them releases CO₂. The ocean exchanges CO₂ with the air in both directions.Atmospherecarbon dioxide, CO2Plants and algaeproducersAnimalsconsumersDead matterand decomposersOceandissolved CO2Fossil fuelscoal, oil, gasPhotosynthesisRespirationEatingDeath and decayExchangeBurialBurning
Photosynthesis
Producers take in CO2 and build it into sugars.
Respiration
Plants, animals and decomposers release CO2 as they break down sugars.
Eating
Carbon passes up food chains.
Death and decay
Decomposers break down dead matter and waste, respiring much of the carbon back to the air.
Ocean exchange
CO2 dissolves into and comes out of seawater.
Burial
Over millions of years, buried remains become coal, oil and gas.
Burning
Burning fossil fuels releases old carbon as CO2 very quickly.

The nitrogen cycle

The nitrogen cycleNitrogen gas in the atmosphere is fixed into ammonium by bacteria such as Rhizobium and by the industrial Haber-Bosch process, and into nitrate by lightning. Soil bacteria turn ammonium into nitrate (nitrification). Plants take up nitrate and build it into proteins and DNA, which pass to animals. Decay returns nitrogen to the soil as ammonium, and denitrifying bacteria turn nitrate back into nitrogen gas.Atmospherenitrogen gas, N2AmmoniumNH4+ in soilNitrateNO3− in soilPlants and animalsproteins and DNAFixation by bacteriaHaber-BoschLightningDenitrificationNitrificationUptake by plantsDecay
Fixation by bacteria
Bacteria such as Rhizobium, in legume root nodules, turn N2 into ammonium.
Lightning
Splits N2; the nitrogen reaches soil as nitrate in rain.
Haber-Bosch
Factories make ammonia for fertilizer from N2 and hydrogen.
Nitrification
Soil bacteria turn ammonium into nitrite, then nitrate.
Uptake by plants
Roots absorb nitrate and ammonium to build proteins and DNA.
Decay
Decomposers return nitrogen from dead matter as ammonium.
Denitrification
Bacteria in low-oxygen soil turn nitrate back into N2.

Nitrogen is often the nutrient that limits plant growth, which is why farmers add fertilizer. Fertilizer that washes off fields into rivers can feed huge blooms of algae. When the algae die and decay, the bacteria breaking them down use up the oxygen in the water, creating low-oxygen "dead zones" such as the one that forms in the Gulf of Mexico most summers.

6. Population growth

With plenty of food and space and nothing holding it back, a population grows exponentially: the more individuals there are, the faster it grows, giving a J-shaped curve. Bacteria in fresh broth grow this way for a while.

No population can keep that up. As it grows, food, space and other resources run short and disease and predators take more. Growth slows and the population levels off near the carrying capacity (K), the largest population the environment can support over time. This S-shaped curve is called logistic growth.

Real populations can overshoot. In 1944, 29 reindeer were released on St. Matthew Island in the Bering Sea, which had no predators. By 1963 there were about 6,000. They ate the island's lichens faster than they could regrow, and after a hard winter only 42 were left.

Limiting factors that get stronger as a population gets denser, like competition for food and disease, are density-dependent. Others, like a hard freeze or a wildfire, hit regardless of density and are density-independent.

Explore: exponential vs logistic

This is a simplified model that uses the standard textbook equations, starting from 10 individuals. Real populations wobble, overshoot and respond to the weather.

Turn on JavaScript to try the sliders. Exponential growth curves upward faster and faster; logistic growth starts the same way, slows as it approaches K and levels off.

7. Living together

RelationshipEffect on each partnerExample
MutualismBoth benefit (+/+)Bees get nectar and pollinate flowers; Rhizobium bacteria get sugar from legumes and give them fixed nitrogen.
CommensalismOne benefits, the other is barely affected (+/0)Barnacles riding on a whale get carried to food-rich water.
ParasitismOne benefits, the other is harmed (+/−)Ticks feeding on a deer; tapeworms in an animal's gut.
CompetitionBoth are harmed (−/−)Mussels and barnacles competing for space on the rock.
PredationPredator benefits, prey is killed (+/−)Sea stars eating mussels; lynx hunting snowshoe hares.

Mutualism, commensalism and parasitism are the three kinds of symbiosis, a close, long-term relationship between two species. The lines between them can blur: a relationship can shift as conditions change.

In the 1930s the Russian biologist Georgii Gause grew two species of Paramecium separately, and each thrived. Grown together on the same food, P. aurelia and P. caudatum could not both last: P. caudatum always died out in his cultures. Yet when he paired P. caudatum with P. bursaria, which fed in a different part of the tube, the two coexisted. This idea, that two species competing for exactly the same resources cannot coexist indefinitely, later became known as the competitive exclusion principle. In nature, competitors often survive by splitting resources, for example by feeding at different times or in different places.

8. Succession

Ecological succession is the gradual change in which species live in a place after a disturbance.

Primary succession starts with no soil at all: on new volcanic rock, or ground left bare by a retreating glacier. Pioneer species such as lichens and mosses arrive first. As they grow, die and help break down rock, a thin soil forms, and grasses, shrubs and eventually trees can follow. This can take centuries. The volcanic island of Surtsey, which rose out of the sea off Iceland in 1963, has been watched since its birth as a natural laboratory of primary succession.

Secondary succession follows a disturbance that removes the community but leaves the soil, such as a fire, a flood or an abandoned farm field. Seeds and roots already in the soil give it a head start, so it is much faster.

Older textbooks describe succession ending in a fixed "climax community." Ecologists now see most communities as always changing, as disturbances of different sizes keep resetting parts of the landscape.

9. Biodiversity and human impacts

Biodiversity is the variety of life, counted at three levels: genetic diversity within a species, the number of species, and the variety of ecosystems. Diverse ecosystems tend to be more stable and provide services people rely on, such as pollination, clean water and fertile soil.

In 2019 the IPBES global assessment, written by hundreds of experts, estimated that around one million species of animals and plants are threatened with extinction. It ranked the main direct causes as:

  1. changes in how land and sea are used, such as clearing forest for farms
  2. direct exploitation: hunting, fishing and logging
  3. climate change
  4. pollution, including fertilizer runoff and plastics
  5. invasive species

Not every trend points down. A 2020 analysis of insect surveys found that land insects declined by roughly 9% per decade on average, while freshwater insects increased in many places, partly thanks to cleaner rivers. Trends differ between places and groups, which is why long-term counts matter. Protected areas, cleaner water and controls on hunting have brought many species back from the brink.

10. How do we know?

  1. 1866 Ernst Haeckel coins the word Oekologie (ecology), from the Greek oikos, "house," for the study of how organisms relate to their surroundings.
  2. 1927 Charles Elton's book Animal Ecology describes food chains and what he called the pyramid of numbers: many small animals at the bottom, few large ones at the top.
  3. 1934 Georgii Gause's The Struggle for Existence reports his Paramecium experiments, the basis for what became the competitive exclusion principle.
  4. 1935 Arthur Tansley coins the word ecosystem for a community together with its physical environment.
  5. 1942 Raymond Lindeman's "trophic-dynamic" paper, based on his study of Cedar Bog Lake in Minnesota, treats an ecosystem as a flow of energy between trophic levels.
  6. 1958 Charles David Keeling begins measuring carbon dioxide at Mauna Loa, Hawaii. His record, the Keeling curve, rose from about 315 parts per million in 1958 to above 420 in the 2020s, with a yearly zigzag as Northern Hemisphere plants grow and shed leaves.
  7. 1963 Gene Likens, F. Herbert Bormann and colleagues begin long-term studies of whole watersheds at the Hubbard Brook Experimental Forest in New Hampshire. When they cleared the forest from one watershed, nitrate in its stream water rose sharply, showing how living plants hold nutrients in an ecosystem.
  8. 1966 Robert Paine publishes his sea star removal experiment, showing that a single predator can keep a community diverse.
  9. 1969 Paine introduces the term keystone species.
  10. 1995 Gray wolves are reintroduced to Yellowstone National Park, starting one of the most closely watched, and most argued-over, natural experiments in ecology.

Quiz: Field Station Check

Talk about it

  1. Paine's experiment removed just one species. Why might it be risky to call a species "unimportant" because it is rare?
  2. Draw a food web for a place near you, such as a park, a pond or a garden. Which species do you think is most likely to be a keystone?
  3. The Yellowstone wolf story is often told as a simple chain of cause and effect. What makes it hard to prove what caused the changes there?
  4. Haber-Bosch fertilizer feeds billions of people but also causes dead zones. How would you weigh those against each other?
  5. Humans now move more carbon and nitrogen than many natural processes do. Does that make people part of these cycles, or something outside them?

Sources

  1. Paine, R. T. (1966). Food web complexity and species diversity. The American Naturalist 100: 65–75. doi:10.1086/282400
  2. Paine, R. T. (1969). A note on trophic complexity and community stability. The American Naturalist 103: 91–93. doi:10.1086/282586
  3. Tansley, A. G. (1935). The use and abuse of vegetational concepts and terms. Ecology 16: 284–307. doi:10.2307/1930070
  4. Lindeman, R. L. (1942). The trophic-dynamic aspect of ecology. Ecology 23: 399–417. doi:10.2307/1930126
  5. Pauly, D. and Christensen, V. (1995). Primary production required to sustain global fisheries. Nature 374: 255–257. doi:10.1038/374255a0
  6. Keeling, C. D. (1960). The concentration and isotopic abundances of carbon dioxide in the atmosphere. Tellus 12: 200–203. doi:10.3402/tellusa.v12i2.9366
  7. Likens, G. E., Bormann, F. H., Johnson, N. M., Fisher, D. W. and Pierce, R. S. (1970). Effects of forest cutting and herbicide treatment on nutrient budgets in the Hubbard Brook watershed-ecosystem. Ecological Monographs 40: 23–47. doi:10.2307/1942440
  8. Ripple, W. J. and Beschta, R. L. (2012). Trophic cascades in Yellowstone: the first 15 years after wolf reintroduction. Biological Conservation 145: 205–213. doi:10.1016/j.biocon.2011.11.005
  9. Kauffman, M. J., Brodie, J. F. and Jules, E. S. (2010). Are wolves saving Yellowstone's aspen? A landscape-level test of a behaviorally mediated trophic cascade. Ecology 91: 2742–2755. doi:10.1890/09-1949.1
  10. Erisman, J. W., Sutton, M. A., Galloway, J., Klimont, Z. and Winiwarter, W. (2008). How a century of ammonia synthesis changed the world. Nature Geoscience 1: 636–639. doi:10.1038/ngeo325
  11. Klein, D. R. (1968). The introduction, increase, and crash of reindeer on St. Matthew Island. The Journal of Wildlife Management 32: 350–367. doi:10.2307/3798981
  12. van Klink, R. et al. (2020). Meta-analysis reveals declines in terrestrial but increases in freshwater insect abundances. Science 368: 417–420. doi:10.1126/science.aax9931
  13. Haeckel, E. (1866). Generelle Morphologie der Organismen. Berlin: Georg Reimer.
  14. Elton, C. (1927). Animal Ecology. London: Sidgwick and Jackson.
  15. Gause, G. F. (1934). The Struggle for Existence. Baltimore: Williams and Wilkins.
  16. IPBES (2019). Global Assessment Report on Biodiversity and Ecosystem Services. ipbes.net/global-assessment
  17. NOAA Global Monitoring Laboratory. Trends in atmospheric carbon dioxide (Mauna Loa). gml.noaa.gov/ccgg/trends
  18. Clark, M. A., Douglas, M. and Choi, J. (2018). Biology 2e, chapters 44–47 (Ecology). OpenStax. openstax.org/books/biology-2e