Chapter 9 DECOMPOSITION

Decomposition is breakdown of chemical bonds formed during the construction of plant and
animal tissue.
It is the end product of the energy pathway begun with photosynthesis.
Decomposition involves respiration: the release of the energy fixed by photosynthesis, C! and
"!.
Decomposition involves several processes: fragmentation, ingestion, egestion, and
concentration.
#ll heterotrophs are decomposers to some extent, but the concept of decomposers is reserved
for those organisms that feed on detritus.
Decomposers include bacteria, fungi and detritivores.
CLASSIFICATION OF DECOMPOSERS
Decomposer bacteria may be aerobic or anaerobic.
#naerobic bacteria use inorganic compounds such as sulfate or nitrate as oxidant that is a
molecule that accepts electrons other than oxygen.
$or example, some bacteria called sulfate reducers can transfer electrons to sulfate
%&'
!(
) reducing it to "!&. ther bacteria, called nitrate reducers, can transfer
electrons to nitrate %*+
(
) reducing it to nitrite %*!
(
). ther nitrate reducers can
reduce nitrate even further to nitrous oxide %*) or nitrogen gas %*!).
*+
(
, !e( , !", -*!( , "!
&ulfate %&'
!(
is reduced to sulfide %&
!(
), typically in the form of hydrogen sulfide
%"!"). .ight electrons are add to sulfate to make sulfide:
acetate , &'
!(
, +"
,
- !C! , "!& , !"!
$ermentation produces organic acids and alcohol.
&ome decomposer bacteria are...
Facultative anaerobes: they use oxygen if available/ if not, they shift to
fermentation.
Obligate anaerobes: they die in the presence of oxygen.
0acteria are the main decomposers of dead animal matter.
$ungi are the main decomposers of plant material.
#fter one group of decomposers has exploited the material to the extent of its ability, a different
group of bacteria and fungi moves in and continue the process of extracting nutrients.
# succession of microflora occurs in the decomposition of organic matter until the material is
finally reduced to inorganic nutrients.
Invertebrate detritivores are involved in the fragmentation of dead organic matter. 1here are four
ma2or groups:
3. Microfauna and icroflora: proto4oans, rotifers and nematodes living in the water film of
soil pores.
!. Meiofauna: animals whose body falls within 3556m ( ! mm, e.g. mites, springtails, some
worms.
+. Macrofauna: between ! mm ( !5 mm long/ e.g. woodlice, spiders, earthworms, beetles,
centipedes, slugs, snails, ants, and harvestmen.
'. Megafauna: over !5 mm, e.g. moles, rabbits, rodents, and earthworms, snails on land,
crabs and mollusks in water.
Microbivores are organisms living in the soil that feed on bacteria, fungi and other small forms.
#mebas, springtails, larvae of beetles, and mites.
7icrobivores control the population of decomposers.
"The invertebrates that feed on detritus are a taxonomically diverse group. In terrestrial
environments, they are usually classified according to their size. This is not an arbitrary basis
for classification, because size dictates their access to resources, as they burrow and crawl
among the cracks and crevices of the litter layer. The microfauna (animals with body widths less
than .! mm" includes protozoans, nematode worms and rotifers. The principle groups of the
mesofauna (body widths between .! mm and # mm" include litter mites ($cari", springtails
(%ollembola", and pot worms (&nchytraeidae". The macrofauna (body widths between # mm to
# mm" and megafauna (body widths greater than # mm" include woodlice (Isopoda",
centipedes (%hilopoda", millipedes ('iplopoda", earthworms ((egadrili", snails and slugs
((ollusca", and the larvae of certain flies ('iptera" and beetles (%oleoptera". The larger animals
are mainly responsible for the initial shredding of plant remains. )oil bacteria and fungi
complete the cycle, consuming the fragmented material. *y their action, detritivores promote a
large+scale redistribution of organic material, and thus contribute directly to the development of
soil structure.8
!tt":##$$$%alban&%e'u#nat$eb#fs'etr%!tl
(eorge Robinson) Assistant Professor) *iological Sciences%
Interesting sites: http:99www.deathonline.net9decomposition9corpse:fauna9index.htm
#nimal decay:
http:99everything!.com9index.pl;node:id<3'=>=39
http:99adipocere.homestead.com9chemistry.html
STA(ES OF DECOMPOSITION
3. Leac!ing is the removal of soluble substances by water, e.g. sugars.
!. Fragentation involves the breaking of the material into smaller pieces/ this may be done
physically or chemically.
1hese two abiotic processes result in the loss of mass and change in the chemical composition
of the detritus.
+. Minerali+ation is the release of organically bound nutrients into an inorganic form available
for plants and microbes. Detritivores oxidi4e organic compounds into smaller and simpler
products.
'. Nutrient iobili+ation is the incorporation of mineral nutrients into microbial biomass.
1hose nutrients not taken up by microbes become available to plants.
In nitrogen minerali4ation, organic nitrogen from decaying plant and animal residues %proteins,
nucleic acids, amino sugars, urea) is converted to ammonia %*"+) and ammonium %*"',) by the
mechanisms listed below. 1his process is also called ammonification. 1he resultant ammonia
can be converted back to organic * %immobili4ation) where it is taken up by microbes and plants
%assimilated), or nitrified to nitrate %*+
(
).
nce *"'
,
has been formed, there are numerous possible fates:
3. It can be assimilated %or immobili4ed) by microbes and plants.
!. It can be held on exchange complexes in soil.
+. It can be fixed in the inner layer portions of clays.
'. It can be volatili4ed as *"+ in decaying vegetation or manure.
?. It can be nitrified by the autotrophic nitrifying bacteria.
T,E PROCESS OF DECOMPOSITION
@iving leaves produce exudates that are washed off by rain and support. 7icroorganisms feed
on the exudates and cells that slough off the surface of the leaf.
nce on the ground, bacteria and fungi attack the leaves. 1he leaves are broken down into
smaller pieces. 1he macro and megafauna are involved in the fragmentation of organic material.
1heir fecal material is rich in microorganisms that continue the chemical decomposition
process.
.arthworms release vitamin 03! into the soil. thers bury and mix organic matter with the
soil particles. .arthworms can process up to 35A of the # hori4on.
In the absence of detritivores, decomposition is slowed down significantly. Bood
decomposition is slowed down by half.
1he soil region immediately surrounding the roots, is known as the r!i+os"!ere and, and the
root surface itself is the r!i+o"lane.
1he rhi4oplane supports a large number of microorganisms.
Coot exudates consist of simple sugars %35), fatty acids %33), amino acids %!3), vitamins %35),
nucleotides %'), and miscellaneous compounds %33). 1he number indicates those identified so
far.
#nimals decomposed faster than plant matter.
&cavengers %crows, foxes, vultures, etc.) consume most of the flesh and some bone material.
Bhat is left is attacked by bacteria rather than fungi.
7aggots can consume a small mammal carcass in about 35 ( 33 days in the summer time.
0acteria and maggots can consume about =5A of the body of an animal. 1he remaining +5A is
mostly hair and bones.
1he fecal matter of large herbivores contains abundance of partially digested organic matter that
provides a rich resource for speciali4ed detritivores in addition to bacteria, fungi and
earthworms.
&ome of the co"ro"!agous 'etritivores are dung beetles and the larva of certain fly species.
Nonhumic substances:
Carbohydrates, proteins, peptides, amino acids, fats, waxes, resins, pigments, and other
low(molecular(weight organic substances.
@abile, easily to be utili4ed and degraded by microorganisms, exibit rapid flux rates/ low
instantaneous concentrations in water
Humic substances %>5A of the organic matter):
"umic acids, fulvic acids, humin/ the formation of humic substances occurs during the
degradation of higher aDuatic and terrestrial plant material %celluloses, hemicelluloses,
and lignin) by fungi and bacteria.
@oss: photolysis, microbial decomposition, aggregation and sedimentation, and
outflows.
"umic acids chelate cations, and help detoxify the soil.
EN-IRONMENTAL CONTROLS ON DECOMPOSERS
1he ma2or factors that influence the rate of decomposition are:
3. Euality of the dead organism as a food source.
!. Fhysical environment, particularly temperature and moisture.
LITTER ./ALIT0
1he nature of the chemical bonds present and the si4e of the three(dimensional structure of the
molecules in which the bonds are formed contribute to the Duality or lack of Duality of the litter.
*ot all carbon compounds are of the same Duality as energy source.
(lucose, simple sugars and soluble proteins are very high Duality sources of carbon and
energy. 1hese compounds decompose very Duickly.
Cellulose and !eicellulose are difficult to break down. 1hey are of moderate Duality, and are
next to being consumed.
Lignins are more complex and variable chemicals. @ignins are a class of chemicals. 1heir long
molecular structure is much folded into complex three(dimensional structures that shield much
of the internal structure from attack by en4ymes
@ignin is a ma2or component of wood and one the slowest components of plant tissue to
decompose.
#s carbohydrates and proteins are removed, the proportion of lignin increases and the
remaining litter becomes of a lesser Duality as energy source.
1he proportion of lignin in the leaves of certain trees makes them less desirable to
decomposers, e.g. oak and pine.
&pecies with less lignin and higher carbohydrate and protein content, support a large
number of detritivores, e. g. aspen, basswood, elm, white ash, and birch.
1he Actinobacteria or #ctinomycetes are a group of Gram(positive bacteria. 7ost are
found in the soil, and they include some of the most common soil life, playing an
important role in decomposition of organic materials, such as cellulose and chitin. 1his
replenishes the supply of nutrients in the soil and is an important part of humus
formation
Pol&"!enols in the litter interfere with the decomposers metabolism and en4yme function
slowing down the process of decomposition.
Pol&"!enols are a group of chemical substances found in plants, characteri4ed by the
presence of more than one phenol group per molecule. 1he polyphenols are responsible
for the coloring of some plants, anthocyanins, e.g. the color of leaves in the autumn.
MEAS/RIN( LITTER *REA1DO2N
1he loss of mass through time is expressed by the following negative exponential function:
3 original ass reaining 4 e
5e6
e < natural logarithm
t < time unit used %years, months, etc.)
k < decomposition coefficient.
1he larger the value of 6, the faster the rate of decomposition and the steeper the curve.
1he most consistent characteristic accounting for variations in the observed rates of decay is
the percentage of the total carbon content that is made up of lignin compounds.
1he greater the amount of carbon tied up in lignin, the lower the consumption rate by
decomposers.
,$ substantial fraction of the mass of the humic acids is in carboxylic acid functional groups,
which endow these molecules with the ability to chelate positively charged multivalent ions
((g--, %a--, .e--, most other "trace elements" of value to plants, as well as other ions that
have no positive biological role, such as %d-- and /b--." This chelation of ions is probably the
most important role of humic acids with respect to living systems. *y chelating the ions, they
facilitate the uptake of these ions by several mechanisms, one of which is preventing their
precipitation0 another seems to be a direct and positive influence on their bioavailability. (any
organisms can exploit dissolved organics to some degree if they are present, and humic acids
may be taken up by this mechanism. $nother paradoxical effect of humic acids is the
detoxification of heavy metals. 1ne might expect them to be made (ore, not less toxic by
humic acids, but the studies that I2ve seen seem to indicate a detoxifying effect.3
Craig 0ingman, &cience .ditor, #$E
http:99www.thekrib.com9Chemistry9humic.html
#dditional information about humic acids: http:99www.phelpstek.com9clients9humic:acid.html
http:99www.ar.wroc.pl9Hweber9humic.htmIstart
http:99aggie(horticulture.tamu.edu9extension9newsletters9hortupdate92un5!9art'2un.html
# study showed that the percentage of nitrogen in absolute terms increased when the mass
decreased during decomposition.
rganisms that decompose organic matter use carbon as a source of energy and nitrogen for building cell structure.
1hey need more carbon than nitrogen. If there is too much carbon, decomposition slows when the nitrogen is used up
and some organisms die. ther organisms form new cell material using their stored nitrogen. In the process more
carbon is burned. 1hus the amount of carbon is reduced while nitrogen is recycled. Decomposition takes longer,
however, when the initial C:N ratio is much above +5.
In the soil, using organic matter with excess carbon can create problems. 1o complete the nitrogen cycle
and continue decomposition, the microbial cells will draw any available soil nitrogen in the proper
proportion to make use of available carbon. 1his is known as 8robbing8 the soil of nitrogen, and delays
availability of nitrogen as a fertili4er for growing plants until some later season when it is no longer being
used in the life(cycles of soil bacteria.
Bhen the energy source, carbon, is less than that reDuired for converting available nitrogen into protein,
organisms make full use of the available carbon and get rid of the excess nitrogen as ammonia. 1his
release of ammonia to the atmosphere produces a loss of nitrogen from the compost pile and should be
kept to a minimum.
# C:N ratio of !5, where C and * are the available Duantities, is the upper limit at which there is no danger of robbing
the soil of nitrogen. If a considerable amount of carbon is in the form of lignin or other resistant materials, the actual
C:N ratio could be larger than !5. 1he C:N ratio is a critical factor in composting to prevent both nitrogen robbing from
the soil and conserving maximum nitrogen in the compost.J
Bashington &tate Kniversity, Bhatcom County
.xtension.http:99whatcom.wsu.edu9ag9compost9fundamentals9needs:carbon:nitrogen.htm
1he optimum ratio in soil organic matter is about 35 carbons to 3 nitrogen, or a C:N ratio of
35:3.
1he nitrogen content reflects the nitrogen remaining in the plant tissue as well as...

In the biomass of the microbes, fungi and bacteria feeding on the plant.
.xcreted products of detritivores.
0acteria and fungi breakdown proteins into amino acids.
L
#mino acids are oxidi4ed into carbon dioxide, water and ammonia.
L
&ome bacteria groups transform ammonium to nitrates.
L
#vailable for plants
1he availability of any particular nutrient to decomposers depends on the ration of energy
supply to nutrient supply, expressed as the carbon:nutrient ratio, C:M
Bhen the C:M ratio of the litter material is high, the supply of that nutrient relative to the energy
source is low.
If the litter does not meet the demand for nutrient, the microbes than obtain the nutrient from
other sources, e.g. soil, and immobili4e it in microbial biomass. 1hese nutrients are then
unavailable to plants.
7inerali4ation and immobili4ation are taking place at the same time. 1he supply rate of mineral
nutrients to the soil during the process of decomposition, the net inerali+ation rate, is the
difference between the rates of minerali4ation and immobili4ation.
In nitrogen immobili4ation, ammonia and nitrate are taken up by microbes and is largely
immobili4ed, or made unavailable to plants, depending on the C:* ratios. Bhen * is abundant,
both microbes and plants assimilate ammonia and nitrate. 1he nitrate undergoes nitrate
reduction by a four(step process and is converted to ammonia by reactions that are similar to
those that occur in denitrification. 1he ammonia is then incorporated into NrebOs cycle
intermediates to form amino acids. $or example, pyruvic acid plus ammonia becomes the amino
acid alanine.
http:99www.ent.iastate.edu9Ipm9Icm9399=9P(!+(399=9nitfert.html
INFL/ENCE OF T,E P,0SICAL EN-IRONMENT
1emperature and moisture influence the rate of microbial activity.
@ow temperature and dry conditions inhibit microbial activity.
7icroflora population and activity is reduced by constant dry conditions and by alternate wetting
and drying.
DECOMPOSITION IN A./ATIC EN-IRONMENTS
Decomposition in aDuatic systems follows the same pattern of leaching, fragmentation,
coloni4ation of detritus by bacteria and fungi, and consumption by invertebrates.
#Duatic fungi attack twigs and leaves in flowing water.
&hredders, aDuatic arthropods, break organic matter into pieces in the process of eating
bacteria and fungi from the surface.
Gra4ers and scrapers feed on algae, bacteria, fungi and organic matter collected on
rocks and large debris.
#lgae take up nutrients and dissolved organic matter from the water.
ther organisms filter particles and fecal matter from the water.
In ponds, lakes and oceans, dead organisms and other organic material, called particulate
organic matter %F7) drift to the bottom.
F7 is ingested, digested and minerali4ed on its way to the bottom. 7uch of the organic matter
settles on the bottom in the form of humic compounds.
I shallow water much of the organic matter arrives in large packages to be digested by benthos
like crabs, snails, and mollusks.
0acteria on the surface sediment can carry aerobic respiration, but a few centimeters below, the
oxygen supply is deplete and anaerobic respiration is used.
*+
(
is used as the electron acceptor instead of oxygen resulting in denitrification, the
conversion of nitrate to molecular nitrogen.
4hen nutrients are not limiting, most phytoplankton has the following molar ratio of
elements5 %565/ 7 !85 !85 !
'enitrification is the microbial reduction of nitrate to di+nitrogen gas. 'enitrification is
sometimes referred to as 2dissimilatory2 nitrate reduction because it occurs in association
with the dissimilation (decomposition" of organic matter.. The denitrification reaction for
the oxidation of organic matter with 9edfield molar proportions is as follows:5
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0acteria can use sulfate and $e
+,
in the middle mud resulting in their reduction.
1he bacteria reDuire: organic matter/ an anoxic environment %or microenvironment)/ and
dissolved sulfate. "ydrogen sulfide gas %"!&) and alkalinity are generated in the process:
&'
!(
, !%C"!) < "!& , !"C+
!(
Dissolved organic matter %D7) in the water column provides a source of fixed carbon for
decomposition.
7acroalgae, phytoplankton and 4ooplankton are a source of D7.
0acteria biomass and large number of phytoplankton are heterotrophic and important in the
recycling of organic matter
# ma2or difference involves dissolved organic matter in the water column, excreted by
phytoplankton and 4ooplankton.
#lgae and phytoplankton excrete large amounts of organic matter during certain stages of their
life cycle, e.g. reproduction.
Bhen phytoplankton and 4ooplankton die, a large amount of organic matter becomes available.
Qooplankton and phytoplankton play a ma2or role in releasing nutrient that is going to be
converted into biomass.
0acteria take up this dissolved organic matter, which is not broken down into inorganic forms,
and convert it into bacterial biomass that in turn becomes food for 4ooplankton.
nly bacteria can assimilate the large amounts of organic matter that is dissolved in water and
convert it into biomass.
Dissolved organic matter is the substrate for the growth of bacteria. D7 and colloidal matter
condense on the surface of air bubbles in the water, forming organic particles on which bacteria
flourish.
Ciliates and 4ooplankton eat bacteria and in turn excrete nutrients in the form of exudates and
fecal pellets in the water.
If there is enough food, 4ooplankton will eat more than it needs and excrete in fecal pellets the
excess ingested food, which will form part of the suspended material.
Microbial loo" is a term used to explain the role of microbes in marine ecosystems.
4hat is the microbial loopA
The microbial loop is a micro+food chain that works within (or along side" the classical food
chain. In the microbial loop the smallest organisms, the heterotrophic bacteria and pico+
plankton use dissolved inorganic material directly (as carbon and energy sources". These
organisms are grazed by flagellates and ciliates. The flagellates and smaller phyto+plankton are
too small to be preyed on directly by copepods, long considered to be a maBor predator of
micro+organisms. Improved detection methods have demonstrated that ciliate organisms
provide an intermediate predator level in the microbial loop. These are then consumed by
copepods and the process continues up the classical food chain.
!tt":##$$$%uib%no#us#aga+ine#u"'ates#Loo"#loo"%!t