Plant litter
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Plant litter (also leaf litter, tree litter, soil litter, litterfall or duff) is dead
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclu ...
material (such as leaves, bark, needles,
twig A twig is a thin, often short, branch of a tree or bush. The buds on the twig are an important diagnostic characteristic, as are the abscission scars where the leaves have fallen away. The color, texture, and patterning of the twig bark ar ...
s, and
cladode Phylloclades and cladodes are flattened, photosynthetic shoots, which are usually considered to be modified branches. The two terms are used either differently or interchangeably by different authors. '' Phyllocladus'', a genus of conifer, is nam ...
s) that have fallen to the ground. This detritus or dead organic material and its constituent nutrients are added to the top layer of soil, commonly known as the litter layer or O horizon ("O" for "organic"). Litter is an important factor in ecosystem dynamics, as it is indicative of ecological productivity and may be useful in predicting regional nutrient cycling and soil fertility.


Characteristics and variability

Litterfall is characterized as fresh, undecomposed, and easily recognizable (by species and type) plant debris. This can be anything from leaves, cones, needles, twigs, bark, seeds/nuts, logs, or reproductive organs (e.g. the stamen of flowering plants). Items larger than 2 cm diameter are referred to as coarse litter, while anything smaller is referred to as fine litter or litter. The type of litterfall is most directly affected by
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...
type. For example, leaf tissues account for about 70 percent of litterfall in forests, but woody litter tends to increase with forest age. In grasslands, there is very little aboveground
perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wid ...
tissue so the annual litterfall is very low and quite nearly equal to the net primary production.Schlesinger, William H. Biogeochemistry: An Analysis of Global Change. 2nd Edition. Academic Press. 108, 135, 152–158, 180–183, 191–194. (1997). In
soil science Soil science is the study of soil as a natural resource on the surface of the Earth including soil formation, classification and mapping; physical, chemical, biological, and fertility properties of soils; and these properties in relation to th ...
, soil litter is classified in three layers, which form on the surface of the O Horizon. These are the L, F, and H layers: The litter layer is quite variable in its thickness, decomposition rate and nutrient content and is affected in part by
seasonality In time series data, seasonality is the presence of variations that occur at specific regular intervals less than a year, such as weekly, monthly, or quarterly. Seasonality may be caused by various factors, such as weather, vacation, and holidays a ...
, plant species, climate, soil fertility, elevation, and
latitude In geography, latitude is a coordinate that specifies the north– south position of a point on the surface of the Earth or another celestial body. Latitude is given as an angle that ranges from –90° at the south pole to 90° at the north pol ...
. The most extreme variability of litterfall is seen as a function of seasonality; each individual species of plant has seasonal losses of certain parts of its body, which can be determined by the collection and classification of plant litterfall throughout the year, and in turn affects the thickness of the litter layer. In tropical environments, the largest amount of debris falls in the latter part of dry seasons and early during wet season. As a result of this variability due to seasons, the decomposition rate for any given area will also be variable. Latitude also has a strong effect on litterfall rates and thickness. Specifically, litterfall declines with increasing latitude. In tropical rainforests, there is a thin litter layer due to the rapid decomposition, while in
boreal forests Taiga (; rus, тайга́, p=tɐjˈɡa; relates to Mongolic and Turkic languages), generally referred to in North America as a boreal forest or snow forest, is a biome characterized by coniferous forests consisting mostly of pines, spruces, ...
, the rate of decomposition is slower and leads to the accumulation of a thick litter layer, also known as a mor. Net primary production works inversely to this trend, suggesting that the accumulation of organic matter is mainly a result of decomposition rate. Surface detritus facilitates the capture and infiltration of rainwater into lower soil layers. Soil litter protects soil aggregates from raindrop impact, preventing the release of clay and silt particles from plugging soil pores. Releasing clay and silt particles reduces the capacity for soil to absorb water and increases cross surface flow, accelerating soil
erosion Erosion is the action of surface processes (such as water flow or wind) that removes soil, rock, or dissolved material from one location on the Earth's crust, and then transports it to another location where it is deposited. Erosion is dis ...
. In addition soil litter reduces wind erosion by preventing soil from losing moisture and providing cover preventing soil transportation. Organic matter accumulation also helps protect soils from
wildfire A wildfire, forest fire, bushfire, wildland fire or rural fire is an unplanned, uncontrolled and unpredictable fire in an area of combustible vegetation. Depending on the type of vegetation present, a wildfire may be more specifically identi ...
damage. Soil litter can be completely removed depending on intensity and severity of wildfires and season. Regions with high frequency wildfires have reduced vegetation density and reduced soil litter accumulation. Climate also influences the depth of plant litter. Typically humid tropical and sub-tropical climates have reduced organic matter layers and horizons due to year-round decomposition and high vegetation density and growth. In temperate and cold climates, litter tends to accumulate and decompose slower due to a shorter growing season.


Net primary productivity

Net
primary production In ecology, primary production is the synthesis of organic compounds from atmospheric or aqueous carbon dioxide. It principally occurs through the process of photosynthesis, which uses light as its source of energy, but it also occurs through ...
and litterfall are intimately connected. In every terrestrial ecosystem, the largest fraction of all net primary production is lost to
herbivores A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
and litter fall. Due to their interconnectedness, global patterns of litterfall are similar to global patterns of net primary productivity. Plant litter, which can be made up of fallen leaves, twigs, seeds, flowers, and other woody debris, makes up a large portion of above ground net primary production of all terrestrial ecosystems.
Fungus A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from t ...
plays a large role in cycling the nutrients from the plant litter back into the ecosystem.


Habitat and food

Litter provides
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
for a variety of organisms.


Plants

Certain plants are specially adapted for germinating and thriving in the litter layers. For example, bluebell ('' Hyacinthoides non-scripta'') shoots puncture the layer to emerge in spring. Some plants with rhizomes, such as common wood sorrel (''
Oxalis acetosella ''Oxalis acetosella'', the wood sorrel or common wood sorrel, is a rhizomatous flowering plant in the family '' Oxalidaceae'', common in most of Europe and parts of Asia. The specific epithet ''acetosella'' refers to its sour taste. The common ...
'') do well in this habitat.


Detritivores and other decomposers

Many organisms that live on the forest floor are
decomposers Decomposers are Organism, organisms that break down dead or decaying organisms; they carry out decomposition, a process possible by only certain kingdoms, such as fungi. Like herbivores and predators, decomposers are heterotrophic, meaning that t ...
, such as
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
. Organisms whose diet consists of plant detritus, such as
earthworm An earthworm is a terrestrial invertebrate that belongs to the phylum Annelida. They exhibit a tube-within-a-tube body plan; they are externally segmented with corresponding internal segmentation; and they usually have setae on all segments. T ...
s, are termed detritivores. The community of decomposers in the litter layer also includes
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
,
amoeba An amoeba (; less commonly spelled ameba or amœba; plural ''am(o)ebas'' or ''am(o)ebae'' ), often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and retracting pseudop ...
, nematodes, rotifer,
tardigrades Tardigrades (), known colloquially as water bears or moss piglets, are a phylum of eight-legged segmented micro-animals. They were first described by the German zoologist Johann August Ephraim Goeze in 1773, who called them Kleiner Wasserbär ...
,
springtail Springtails (Collembola) form the largest of the three lineages of modern hexapods that are no longer considered insects (the other two are the Protura and Diplura). Although the three orders are sometimes grouped together in a class called Ento ...
s, cryptostigmata, potworms, insect
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
e, mollusks, oribatid mites,
woodlice A woodlouse (plural woodlice) is an isopod crustacean from the polyphyleticThe current consensus is that Oniscidea is actually triphyletic suborder Oniscidea within the order Isopoda. They get their name from often being found in old wood ...
, and
millipede Millipedes are a group of arthropods that are characterised by having two pairs of jointed legs on most body segments; they are known scientifically as the class Diplopoda, the name derived from this feature. Each double-legged segment is a resu ...
s. Even some species of microcrustaceans, especially copepods (for instance '' Bryocyclops spp''., ''Graeteriella spp.'',''Olmeccyclops hondo'', ''Moraria spp''.,''Bryocamptus spp''., ''Atheyella spp''.) live in moist leaf litter habitats and play an important role as predators and decomposers. The consumption of the litterfall by decomposers results in the breakdown of simple carbon compounds into
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is trans ...
(CO2) and
water Water (chemical formula ) is an Inorganic compound, inorganic, transparent, tasteless, odorless, and Color of water, nearly colorless chemical substance, which is the main constituent of Earth's hydrosphere and the fluids of all known living ...
(H2O), and releases inorganic
ions An ion () is an atom or molecule with a net electrical charge. The charge of an electron is considered to be negative by convention and this charge is equal and opposite to the charge of a proton, which is considered to be positive by conven ...
(like
nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...
and
phosphorus Phosphorus is a chemical element with the symbol P and atomic number 15. Elemental phosphorus exists in two major forms, white phosphorus and red phosphorus, but because it is highly reactive, phosphorus is never found as a free element on Ear ...
) into the soil where the surrounding plants can then reabsorb the nutrients that were shed as litterfall. In this way, litterfall becomes an important part of the nutrient cycle that sustains forest environments. As litter decomposes, nutrients are released into the environment. The portion of the litter that is not readily decomposable is known as humus. Litter aids in soil moisture retention by cooling the ground surface and holding moisture in decaying organic matter. The flora and fauna working to decompose soil litter also aid in soil respiration. A litter layer of decomposing biomass provides a continuous energy source for macro- and micro-organisms.


Larger animals

Numerous reptiles, amphibians,
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s, and even some mammals rely on litter for shelter and forage. Amphibians such as
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All t ...
s and
caecilian Caecilians (; ) are a group of limbless, vermiform or serpentine amphibians. They mostly live hidden in the ground and in stream substrates, making them the least familiar order of amphibians. Caecilians are mostly distributed in the tropics o ...
s inhabit the damp
microclimate A microclimate (or micro-climate) is a local set of atmospheric conditions that differ from those in the surrounding areas, often with a slight difference but sometimes with a substantial one. The term may refer to areas as small as a few squ ...
underneath fallen leaves for part or all of their life cycle. This makes them difficult to observe. A
BBC #REDIRECT BBC #REDIRECT BBC Here i going to introduce about the best teacher of my life b BALAJI sir. He is the precious gift that I got befor 2yrs . How has helped and thought all the concept and made my success in the 10th board exam. ...
...
film crew captured footage of a female caecilian with young for the first time in a documentary that aired in 2008. Some species of birds, such as the
ovenbird The ovenbird (''Seiurus aurocapilla'') is a small songbird of the New World warbler family (Parulidae). This migratory bird breeds in eastern North America and winters in Central America, many Caribbean islands, Florida and northern Venezuela. ...
of eastern North America for example, require leaf litter for both foraging and material for
nest A nest is a structure built for certain animals to hold eggs or young. Although nests are most closely associated with birds, members of all classes of vertebrates and some invertebrates construct nests. They may be composed of organic materi ...
s. Sometimes litterfall even provides energy to much larger mammals, such as in
boreal forest Taiga (; rus, тайга́, p=tɐjˈɡa; relates to Mongolic and Turkic languages), generally referred to in North America as a boreal forest or snow forest, is a biome characterized by coniferous forests consisting mostly of pines, spruc ...
s where lichen litterfall is one of the main constituents of wintering
deer Deer or true deer are hoofed ruminant mammals forming the family Cervidae. The two main groups of deer are the Cervinae, including the muntjac, the elk (wapiti), the red deer, and the fallow deer; and the Capreolinae, including the re ...
and elk diets.


Nutrient cycle

During leaf senescence, a portion of the plant's nutrients are reabsorbed from the leaves. The nutrient concentrations in litterfall differ from the nutrient concentrations in the mature foliage by the reabsorption of constituents during leaf senescence. Plants that grow in areas with low nutrient availability tend to produce litter with low nutrient concentrations, as a larger proportion of the available nutrients is reabsorbed. After senescence, the nutrient-enriched leaves become litterfall and settle on the soil below. Litterfall is the dominant pathway for nutrient return to the soil, especially for
nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...
(N) and
phosphorus Phosphorus is a chemical element with the symbol P and atomic number 15. Elemental phosphorus exists in two major forms, white phosphorus and red phosphorus, but because it is highly reactive, phosphorus is never found as a free element on Ear ...
(P). The accumulation of these nutrients in the top layer of soil is known as soil immobilization. Once the litterfall has settled, decomposition of the litter layer, accomplished through the leaching of nutrients by rainfall and throughfall and by the efforts of detritivores, releases the breakdown products into the soil below and therefore contributes to the cation exchange capacity of the soil. This holds especially true for highly weathered tropical soils.
Leaching Leaching is the loss or extraction of certain materials from a carrier into a liquid (usually, but not always a solvent). and may refer to: * Leaching (agriculture), the loss of water-soluble plant nutrients from the soil; or applying a small amou ...
is the process by which cations such as
iron Iron () is a chemical element with Symbol (chemistry), symbol Fe (from la, Wikt:ferrum, ferrum) and atomic number 26. It is a metal that belongs to the first transition series and group 8 element, group 8 of the periodic table. It is, Abundanc ...
(Fe) and
aluminum Aluminium (aluminum in American and Canadian English) is a chemical element with the symbol Al and atomic number 13. Aluminium has a density lower than those of other common metals, at approximately one third that of steel. It ha ...
(Al), as well as organic matter are removed from the litterfall and transported downward into the soil below. This process is known as
podzolization Podsolisation is an extreme form of leaching which causes the eluviation of iron and aluminium sesquioxides. The process generally occurs in areas where precipitation is greater than evapotranspiration. The minerals are removed by a process kno ...
and is particularly intense in boreal and cool temperate forests that are mainly constituted by coniferous pines whose litterfall is rich in phenolic compounds and
fulvic acid Humic substances (HS) are organic compounds that are important components of humus, the major organic fraction of soil, peat, and coal (and also a constituent of many upland streams, dystrophic lakes, and ocean water). For a long era in the 19th an ...
. By the process of biological decomposition by microfauna, bacteria, and fungi, CO2 and H2O, nutrient elements, and a decomposition-resistant organic substance called humus are released. Humus composes the bulk of organic matter in the lower soil profile. The decline of nutrient ratios is also a function of decomposition of litterfall (i.e. as litterfall decomposes, more nutrients enter the soil below and the litter will have a lower nutrient ratio). Litterfall containing high nutrient concentrations will decompose more rapidly and
asymptote In analytic geometry, an asymptote () of a curve is a line such that the distance between the curve and the line approaches zero as one or both of the ''x'' or ''y'' coordinates tends to infinity. In projective geometry and related context ...
as those nutrients decrease. Knowing this, ecologists have been able to use nutrient concentrations as measured by
remote sensing Remote sensing is the acquisition of information about an object or phenomenon without making physical contact with the object, in contrast to in situ or on-site observation. The term is applied especially to acquiring information about Eart ...
as an index of a potential rate of decomposition for any given area. Globally, data from various forest ecosystems shows an inverse relationship in the decline in nutrient ratios to the apparent nutrition availability of the forest. Once nutrients have re-entered the soil, the plants can then reabsorb them through their
root In vascular plants, the roots are the organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often below the su ...
s. Therefore, nutrient reabsorption during senescence presents an opportunity for a plant's future net primary production use. A relationship between nutrient stores can also be defined as: :annual storage of nutrients in plant tissues + replacement of losses from litterfall and leaching = the amount of uptake in an ecosystem


Ocean litter

Non-terrestrial litterfall follows a very different path. Litter is produced both inland by
terrestrial plant A terrestrial plant is a plant that grows on, in, or from land. Other types of plants are aquatic (living in water), epiphytic (living on trees) and lithophytic (living in or on rocks). The distinction between aquatic and terrestrial plants i ...
s and moved to the coast by fluvial processes, and by mangrove ecosystems. From the coast Robertson & Daniel 1989 found it is then removed by the
tide Tides are the rise and fall of sea levels caused by the combined effects of the gravitational forces exerted by the Moon (and to a much lesser extent, the Sun) and are also caused by the Earth and Moon orbiting one another. Tide tables ...
, crabs and microbes. They also noticed that which of those three is most significant depends on the tidal regime. Nordhaus et al. 2011 find crabs forage for leaves at low tide and if their detritivory is the predominant disposal route, they can take 80% of leaf material. Bakkar et al 2017 studied the chemical contribution of the resulting crab defecation. They find crabs pass a noticeable amount of undegraded lignins to both the
sediment Sediment is a naturally occurring material that is broken down by processes of weathering and erosion, and is subsequently transported by the action of wind, water, or ice or by the force of gravity acting on the particles. For example, sa ...
s and water composition. They also find that the exact carbonaceous contribution of each plant species can be traced from the plant, through the crab, to its sediment or water disposition in this way. Crabs are usually the only significant macrofauna in this process, however Raw et al 2017 find ''
Terebralia palustris ''Terebralia palustris'', common name the giant mangrove whelk, is a species of brackish-water snail, a gastropod mollusk in the family Potamididae. This tropical species which inhabits mangrove environments of the Indo-West Pacific region,Houbr ...
'' competes with crabs unusually vigorously in
southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical south-eastern region of Asia, consisting of the regions that are situated south of mainlan ...
.


Collection and analysis

The main objectives of litterfall sampling and analysis are to quantify litterfall production and chemical composition over time in order to assess the variation in litterfall quantities, and hence its role in nutrient cycling across an environmental gradient of
climate Climate is the long-term weather pattern in an area, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteorologi ...
(moisture and temperature) and soil conditions. Ecologists employ a simple approach to the collection of litterfall, most of which centers around one piece of equipment, known as a litterbag. A litterbag is simply any type of container that can be set out in any given area for a specified amount of time to collect the plant litter that falls from the
canopy Canopy may refer to: Plants * Canopy (biology), aboveground portion of plant community or crop (including forests) * Canopy (grape), aboveground portion of grapes Religion and ceremonies * Baldachin or canopy of state, typically placed over an ...
above. Litterbags are generally set in random locations within a given area and marked with
GPS The Global Positioning System (GPS), originally Navstar GPS, is a satellite-based radionavigation system owned by the United States government and operated by the United States Space Force. It is one of the global navigation satellite sy ...
or local coordinates, and then monitored on a specific time interval. Once the samples have been collected, they are usually classified on type, size and
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
(if possible) and recorded on a spreadsheet. When measuring bulk litterfall for an area, ecologists will weigh the dry contents of the litterbag. By this method litterfall flux can be defined as: :litterfall (kg m−2 yr−1) = total litter mass (kg) / litterbag area (m2) The litterbag may also be used to study decomposition of the litter layer. By confining fresh litter in the mesh bags and placing them on the ground, an ecologist can monitor and collect the decay measurements of that litter. An exponential decay pattern has been produced by this type of experiment: \frac = e^, where X_o is the initial leaf litter and k is a constant fraction of detrital mass. The mass-balance approach is also utilized in these experiments and suggests that the decomposition for a given amount of time should equal the input of litterfall for that same amount of time. :litterfall = ''k''(detrital mass) For study various groups from edaphic fauna you need a different mesh sizes in the litterbags


Issues


Change due to invasive earthworms

In some regions of glaciated North America, earthworms have been introduced where they are not native. Non-native earthworms have led to environmental changes by accelerating the rate of decomposition of litter. These changes are being studied, but may have negative impacts on some inhabitants such as salamanders.


Forest litter raking

Leaf litter accumulation depends on factors like wind, decomposition rate and species composition of the forest. The quantity, depth and humidity of leaf litter varies in different habitats. The leaf litter found in primary forests is more abundant, deeper and holds more humidity than in secondary forests. This condition also allows for a more stable leaf litter quantity throughout the year. This thin, delicate layer of organic material can be easily affected by humans. For instance, forest litter raking as a replacement for straw in husbandry is an old non-timber practice in forest management that has been widespread in Europe since the seventeenth century. In 1853, an estimated 50 Tg of dry litter per year was raked in European forests, when the practice reached its peak. This human disturbance, if not combined with other degradation factors, could promote podzolisation; if managed properly (for example, by burying litter removed after its use in animal husbandry), even the repeated removal of forest biomass may not have negative effects on
pedogenesis Soil formation, also known as pedogenesis, is the process of soil genesis as regulated by the effects of place, environment, and history. Biogeochemical processes act to both create and destroy order ( anisotropy) within soils. These alterations ...
.


See also

*
Coarse woody debris Coarse woody debris (CWD) or coarse woody habitat (CWH) refers to fallen dead trees and the remains of large branches on the ground in forests and in rivers or wetlands.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). C ...
* Detritus *
Forest floor The forest floor, also called detritus, duff and the O horizon, is one of the most distinctive features of a forest ecosystem. It mainly consists of shed vegetative parts, such as leaves, branches, bark, and stems, existing in various stages ...
* Leaf litter sieve *
Leaf mold Leaf mold (spelled leaf mould outside of the United States) is the compost produced by decomposition of shaded deciduous shrub and tree leaves, primarily by fungal breakdown in a slower cooler manner as opposed to the bacterial degradation of lea ...
(a type of compost) * Soil horizon


References


External links


forestresearch.gov.uk
{{Authority control Biology terminology Ecological restoration Ecology terminology Ecology Environmental terminology Habitat Soil improvers