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s water moves through dead organic matter it dissolves and carries with it the water soluble components these are then taken up by organisms in the soil react with mineral soil or are transported beyond the confines of the ecosystem and are considered lost to it 20 271 280 newly shed leaves and newly dead animals have high concentrations of water soluble components and include sugars amino acids and mineral nutrients leaching is more important in wet environments and less important in dry ones 10 69 77 fragmentation processes break organic material into smaller pieces exposing new surfaces for colonization by microbes freshly shed leaf litter may be inaccessible due to an outer layer of cuticle or bark and cell contents are protected by a cell wall newly dead animals may be covered by an exoskeleton fragmentation processes which break through these protective layers accelerate the rate of microbial decomposition 19 184 animals fragment detritus as they hunt for food as does passage through the gut freeze thaw cycles and cycles of wetting and drying also fragment dead material 19 186 the chemical alteration of the dead organic matter is primarily achieved through bacterial and fungal action fungal hyphae produce enzymes that can break through the tough outer structures surrounding dead plant material they also produce enzymes that break down lignin which allows them access to both cell contents and the nitrogen in the lignin fungi can transfer carbon and nitrogen through their hyphal networks and thus unlike bacteria are not dependent solely on locally available resources 19 186 decomposition rates decomposition rates vary among ecosystems 21 the rate of decomposition is governed by three sets of factors the physical environment temperature moisture and soil properties the quantity and quality of the dead material available to decomposers and the nature of the microbial community itself 19 194 temperature controls the rate of microbial respiration the higher the temperature the faster the microbial decomposition occurs temperature also affects soil moisture which affects decomposition freeze thaw cycles also affect decomposition freezing temperatures kill soil microorganisms which allows leaching to play a more important role in moving nutrients around this can be especially important as the soil thaws in the spring creating a pulse of nutrients that become available 20 280 decomposition rates are low under very wet or very dry conditions decomposition rates are highest in wet moist conditions with adequate levels of oxygen wet soils tend to become deficient in oxygen this is especially true in wetlands which slows microbial growth in dry soils decomposition slows as well but bacteria continue to grow albeit at a slower rate even after soils become too dry to support plant growth 19 200 dynamics and resilience further information resistance ecology and ecological resilience ecosystems are dynamic entities they are subject to periodic disturbances and are always in the process of recovering from past disturbances 22 347 when a perturbation occurs an ecosystem responds by moving away from its initial state the tendency of an ecosystem to remain close to its equilibrium state despite that disturbance is termed its resistance the capacity of a system to absorb disturbance and reorganize while undergoing change so as to retain essentially the same function structure identity and feedbacks is termed its ecological resilience 23 24 resilience thinking also includes humanity as an integral part of the biosphere where we are dependent on ecosystem services for our survival and must build and maintain their natural capacities to withstand shocks and disturbances 25 time plays a central role over a wide range for example in the slow development of soil from bare rock and the faster recovery of a community from disturbance 14 67 disturbance also plays an important role in ecological processes f stuart chapin and coauthors define disturbance as a relatively discrete event in time that removes plant biomass 22 346 this can range from herbivore outbreaks treefalls fires hurricanes floods glacial advances to volcanic eruptions such disturbances can cause large changes in plant animal and microbe populations as well as soil organic matter content disturbance is followed by succession a directional change in ecosystem structure and functioning resulting from biotically driven changes in resource supply 2 470 the frequency and severity of disturbance determine the way it affects ecosystem function a major disturbance like a volcanic eruption or glacial advance and retreat leave behind soils that lack plants animals or organic matter ecosystems that experience such disturbances undergo primary succession a less severe disturbance like forest fires hurricanes or cultivation result in secondary succession and a faster recovery 22 348 more severe and more frequent disturbance result in longer recovery times from one year to another ecosystems experience variation in their biotic and abiotic environments a drought a colder than usual winter and a pest outbreak all are short term variability in environmental conditions animal populations vary from year to year building up during resource rich periods and crashing as they overshoot their food supply longer term changes also shape ecosystem processes for example the forests of eastern north america still show legacies of cultivation which ceased in 1850 when large areas were reverted to forests 22 340 another example is the methane production in eastern siberian lakes that is controlled by organic matter which accumulated during the pleistocene 26 a freshwater lake in gran canaria an island of the canary islands clear boundaries make lakes convenient to study using an ecosystem approach nutrient cycling see also nutrient cycle biogeochemical cycle and nitrogen cycle biological nitrogen cycling ecosystems continually exchange energy and carbon with the wider environment mineral nutrients on the other hand are mostly cycled back and forth between plants animals microbes and the soil most nitrogen enters ecosystems through biological nitrogen fixation is deposited through precipitation dust gases or is applied as fertilizer 20 266 most terrestrial ecosystems are nitrogen limited in the short term making nitrogen cycling an important control on ecosystem production 20 289 over the long term phosphorus availability can also be critical 27 macronutrients which are required by all plants in large quantities include the primary nutrients which are most limiting as they are used in largest amounts nitrogen phosphorus potassium 28 231 secondary major nutrients less often limiting include calcium magnesium sulfur micronutrients required by all plants in small quantities include boron chloride copper iron manganese molybdenum zinc finally there are also beneficial nutrients which may be required by certain plants or by plants under specific environmental conditions aluminum cobalt iodine nickel selenium silicon sodium vanadium 28 231 until modern times nitrogen fixation was the major source of nitrogen for ecosystems nitrogen fixing bacteria either live symbiotically with plants or live freely in the soil the energetic cost is high for plants that support nitrogen fixing symbionts as much as 25 of gross primary production when measured in controlled conditions many members of the legume plant family support nitrogen fixing symbionts some cyanobacteria are also capable of nitrogen fixation these are phototrophs which carry out photosynthesis like other nitrogen fixing bacteria they can either be free living or have symbiotic relationships with plants 22 360 other sources of nitrogen include acid deposition produced through the combustion of fossil fuels ammonia gas which evaporates from agricultural fields which have had fertilizers applied to them and dust 20 270 anthropogenic nitrogen inputs account for about 80 of all nitrogen fluxes in ecosystems 20 270 when plant tissues are shed or are eaten the nitrogen in those tissues becomes available to animals and microbes microbial decomposition releases nitrogen compounds from dead organic matter in the soil where plants fungi and bacteria compete for it some soil bacteria use organic nitrogen containing compounds as a source of carbon and release ammonium ions into the soil this process is known as nitrogen mineralization others convert ammonium to nitrite and nitrate ions a process known as nitrification nitric oxide and nitrous oxide are also produced during nitrification 20 277 under nitrogen rich and oxygen poor conditions nitrates and nitrites are converted to nitrogen gas a process known as denitrification 20 281 mycorrhizal fungi which are symbiotic with plant roots use carbohydrates supplied by the plants and in return transfer phosphorus and nitrogen compounds back to the plant roots 29 30 this is an important pathway of organic nitrogen transfer from dead organic matter to plants this mechanism may contribute to more than 70 tg of annually assimilated plant nitrogen thereby playing a critical role in global nutrient cycling and ecosystem function 30 phosphorus enters ecosystems through weathering as ecosystems age this supply diminishes making phosphorus limitation more common in older landscapes especially in the tropics 20 287 290 calcium and sulfur are also produced by weathering but acid deposition is an important source of sulfur in many ecosystems although magnesium and manganese are produced by weathering exchanges between soil organic matter and living cells account for a significant portion of ecosystem fluxes potassium is primarily cycled between living cells and soil organic matter 20 291 function and biodiversity main article biodiversity see also ecosystem diversity loch lomond in scotland forms a relatively isolated ecosystem the fish community of this lake has remained stable over a long period until a number of introductions in the 1970s restructured its food web 31 spiny forest at ifaty madagascar featuring various adansonia baobab species alluaudia procera madagascar ocotillo and other vegetation biodiversity plays an important role in ecosystem functioning 32 449 453 ecosystem processes are driven by the species in an ecosystem the nature of the individual species and the relative abundance of organisms among these species ecosystem processes are the net effect of the actions of individual organisms as they interact with their environment ecological theory suggests that in order to coexist species must have some level of limiting similarity they must be different from one another in some fundamental way otherwise one species would competitively exclude the other 33 despite this the cumulative effect of additional species in an ecosystem is not linear additional species may enhance nitrogen retention for example however beyond some level of species richness 11 331 additional species may have little additive effect unless they differ substantially from species already present 11 324 this is the case for example for exotic species 11 321 the addition or loss of species that are ecologically similar to those already present in an ecosystem tends to only have a small effect on ecosystem function ecologically distinct species on the other hand have a much larger effect similarly dominant species have a large effect on ecosystem function while rare species tend to have a small effect keystone species tend to have an effect on ecosystem function that is disproportionate to their abundance in an ecosystem 11 324 an ecosystem engineer is any organism that creates significantly modifies maintains or destroys a habitat 34 study approaches ecosystem ecology main article ecosystem ecology see also ecosystem model a hydrothermal vent is an ecosystem on the ocean floor the scale bar is 1 m ecosystem ecology is the study of the interactions between organisms and their environment as an integrated system 2 458 the size of ecosystems can range up to ten orders of magnitude from the surface layers of rocks to the surface of the planet 4 6 the hubbard brook ecosystem study started in 1963 to study the white mountains in new hampshire it was the first successful attempt to study an entire watershed as an ecosystem the study used stream chemistry as a means of monitoring ecosystem properties and developed a detailed biogeochemical model of the ecosystem 35 long term research at the site led to the discovery of acid rain in north america in 1972 researchers documented the depletion of soil cations especially calcium over the next several decades 36 ecosystems can be studied through a variety of approaches theoretical studies studies monitoring specific ecosystems over long periods of time those that look at differences between ecosystems to elucidate how they work and direct manipulative experimentation 37 studies can be carried out at a variety of scales ranging from whole ecosystem studies to studying microcosms or mesocosms simplified representations of ecosystems 38 american ecologist stephen r carpenter has argued that microcosm experiments can be irrelevant and diversionary if they are not carried out in conjunction with field studies done at the ecosystem scale in such cases microcosm experiments may fail to accurately predict ecosystem level dynamics 39 classifications further information ecosystem classification and biogeoclimatic ecosystem classification biomes are general classes or categories of ecosystems 4 14 however there is no clear distinction between biomes and ecosystems 40 biomes are always defined at a very general level ecosystems can be described at levels that range from very general in which case the names are sometimes the same as those of biomes to very specific such as wet coastal needle leafed forests biomes vary due to global variations in climate biomes are often defined by their structure at a general level for example tropical forests temperate grasslands and arctic tundra 4 14 there can be any degree of subcategories among ecosystem types that comprise a biome e g needle leafed boreal forests or wet tropical forests although ecosystems are most commonly categorized by their structure and geography there are also other ways to categorize and classify ecosystems such as by their level of human impact see anthropogenic biome or by their integration with social processes or technological processes or their novelty e g novel ecosystem each of these taxonomies of ecosystems tends to emphasize different structural or functional properties 41 none of these is the best classification ecosystem classifications are specific kinds of ecological classifications that consider all four elements of the definition of ecosystems a biotic component an abiotic complex the interactions between and within them and the physical space they occupy 41 different approaches to ecological classifications have been developed in terrestrial fres...
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