Академический Документы
Профессиональный Документы
Культура Документы
I. General Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
A. Decomposition, Nutrient Turnover, and Global
Climate Change . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
B. Biomass Distribution between Soil and Above‐Ground
Ecosystem Compartments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
C. The Importance of Balance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
I. GENERAL REMARKS
Very few people without some ecological background turn their attention
to dead organic matter and its decay. The reason is simple: the processes
on which this book focuses occur, to some extent in an ‘‘invisible’’ way,
without such spectacular events as blooming flowers, singing birds, or color-
ful butterflies. What more easily attracts our attention is the opposite side
of the organic matter turnover: the production. The importance of organic
matter production seems obvious to everybody, not only to specialists—this
is the source of our crops and fodder for animals which are, in turn, utilized
as food for humans; this is the timber used for housing, furniture, and paper
production. The list can easily be made much longer. Life is production, and
production means the synthesis of organic compounds from inorganic
chemical elements. Nevertheless, those of us closer to agriculture or forestry
are perfectly familiar also with the opposite side of the story—organic
matter decomposition. For centuries, well before the development of modern
science, farmers knew that in order to sustain agricultural production for
years to come, their fields must be supplemented with nutrients. Agricultural
fields are fertilized with manure, which undergoes the natural process of
decomposition, eventually leading to the release of mineral nutrients in-
dispensable for plants to grow. Some agriculture practices show that
farmers have known that fertilization with organic manure is not the goal
by itself—yearly burning of stubble on meadows, still a common practice in
many parts of the world, reveals recognition of the necessity of mineraliza-
tion of organic matter. The burning of organic residues dramatically short-
ens the time needed for release of nutrients and supplements soil with
mineral nutrients, which can be easily utilized by plants.
These two equations summarize the initial synthesis and the final mineral-
ization. The enormous set of processes is much more complicated, of course,
with an overwhelming variety of organic compounds produced by plants
from a range of inorganic compounds and mineral nutrients, transformed
and complicated even further by consumers. The chemical composition of
litter—the substrate for decomposition processes—is described in detail in
Chapter 2.
Decomposition undergoes a number of steps, leading from complicated
organic compounds through simpler compounds to mineral nutrients, and,
under certain circumstances, not all chemical elements return to their original
inorganic form (Chapter 4). Actually, under the common term ‘‘decomposi-
tion,’’ most scientists understand a whole set of biochemical/microbial
processes, even those opposite to the strict meaning of the term, such as
polymerization of long chains of secondary organic matter collectively called
‘‘humus.’’ However, such processes, going in a direction opposite to actual
degradation, rely on substrates released by earlier partial decay of primary
organic matter. In that sense, they belong to the long list of complicated
processes of dead organic matter transformations and cannot be considered
separately from strict decomposition (cf. Chapter 6). These processes would
be impossible without the billions of microorganisms per gram soil, either
directly engaged in microbial enzymatic degradation of dead organic matter
or indirectly aVecting these processes. The taxonomy of soil organisms,
belonging to such divergent groups as bacteria, fungi, protozoans, potworms,
earthworms, insects, and even vertebrates, exceeds the scope of this book.
However, our feeling is that the book would be incomplete without at least
a short introduction to soil ecology and a presentation of the principal
INTRODUCTION 3
Figure 1 Natural ecosystems have relatively closed internal cycles of most nutrients,
with only minor exchange with the environment outside the ecosystem, such as input
with precipitation and output with water; in some ecosystems, aerial weathering may
be also important.
People with little knowledge of soil biology tend to notice only aboveground
life, manifested by an amazing richness of plants and animals. However,
most heterotrophic life is tied to the soil. Considering the biomass of the most
common groups of terrestrial heterotrophs (animals and microorganisms), it
appears that those animals which most people consider the most abundant
and, possibly, most important for ecosystem function, are, in fact, negligible
in comparison to the ground‐living and soil‐dwelling ones (Table 1). One of
the most spectacular examples among those given in Table 1 are earthworms,
which, in certain agricultural soils, can reach a biomass of up to two tons per
hectare. There is no group of aboveground animals that compares to earth-
worms. The comparison is even more striking for microorganisms, such as
bacteria and fungi—the two groups responsible for most of the organic
matter decomposition in soil. Moreover, the distribution of live biomass
between soil and the aboveground ecosystem compartments illustrates the
importance of decomposers to a limited extent only, because the actual
energy flow through any trophic level is proportional not to the biomass
itself (the ‘‘standing crop’’, [Sc]) but to its total production per time unit (e.g.,
a year). This, in turn, is the product of the standing crop and the rotation
time , the value indicating how many times a year the biomass of a certain
group of organisms (e.g., a population) is produced. The rotation time is the
reciprocal of the average life span t of an individual in a population: y ¼ 1t
where t is given in years. Then, the yearly production is P ¼ Sc y. This
simple equation has far‐reaching consequences for explaining the relative
importance of decomposers in an ecosystem. As these are mostly microor-
ganisms and invertebrates with very high rotation times (especially the
former), their eVect on energy transfer is a few orders of magnitude higher
than would result directly from their biomass. Because, as has been men-
tioned, the energy transfer is linked directly to carbon oxidation (‘‘respira-
tion’’); also, CO2 production by soil organisms is much higher than would
be expected from their biomass alone, making these particular groups of
10 BJÖRN BERG AND RYSZARD LASKOWSKI
Figure 3 Two extreme and contradictory but still possible scenarios for the eVect of
increased atmospheric CO2 concentration on soil organic matter: a positive feedback
loop leads to even further increase in CO2 concentration and global temperature; a
negative feedback mechanism counterbalances the eVect of global warming through
increased carbon sequestration in soil organic matter.
So far, very few detailed studies have been done on biomass and nutrient
distribution among diVerent compartments of forest ecosystems because of
the extreme laboriousness of such research. One notable exception is an
extensive study done in selected mixed forests in Belgium in the 1960s,
which was summarized by Duvigneaud and Denaeyer‐De Smet (1970). The
research team measured and calculated virtually every detail of the biogeo-
chemical cycles in the forests, giving an unmatched body of data on biomass
and dead organic matter distribution in the ecosystems, uptake of nutrients
from soil, their retention in plants, and their return to forest floor with litter
fall. The results of such studies clearly stress the importance of soil organic
matter deposits and mineralization. For example, in the forest presented by
Duvigneaud and Denaeyer‐De Smet (1970), the total aboveground plant
biomass was estimated at 121 t ha1, which together with belowground
biomass of 35 t ha1 (plant roots) gave 156 t of live plant organic matter
biomass per hectare. These researchers were among the first who noticed
that the soil organic matter (SOM) pool was larger than that of aboveground
biomass and not much lower than the total plant biomass in that forest: it
was estimated to 125 t ha1 plus approximately 4.8 t ha1 accumulated on
the soil surface as plant litter—the most easily degradable pool of dead
organic matter. Thus, in temperate hardwoods similar to those studied by
the Belgian group, we may expect that approximately as much organic
12 BJÖRN BERG AND RYSZARD LASKOWSKI
Figure 4 Main organic matter pools in a typical temperate forest ecosystem: live
biomass (aboveground and underground) and dead organic matter (forest litter and
soil organic matter). Data from Duvigneaud and Denaeyer‐De Smet, 1970.
‘‘Balance’’ is an often used term in ecology studies and the term is also
used with respect to humus layers and nutrients stored in humus. There are
numerous articles using synonymous terms, such as ‘‘steady state,’’ which,
Figure 5 (continued )
INTRODUCTION 15
for the amount of stored humus, is assumed to reflect a balance between the
production of litter and the decomposition. In other words, the amount of
16 BJÖRN BERG AND RYSZARD LASKOWSKI
long run because plants adapted to poorer soil conditions are not able to use
all available soluble mobile nutrients in a short time, and their excess can be
irreparably leached from the ecosystem. The extreme example of such quick
leaching of nutrients from an ecosystem can be events such as wildfires,
already mentioned, and human‐made fires, such as those still used in many
countries to ‘‘fertilize’’ meadows in the spring.
The important message that emerges from these considerations is that the
existence of any natural ecosystem depends, to a large extent, on the balance
between the release rate of nutrients from decomposing organic matter and
the rate of their uptake by plants. Specifically, this also means that ecosystems
diVer not only from the structural point of view, such as species composition,
but also functionally. As well as an untrained person can distinguish a pine
forest from an alder wood with the naked eye, an ecologist can recognize
them by looking at their productivity, nutrient pools, and fluxes.
This, together with the biomass distribution in a forest presented on pre-
ceding pages, clearly underlines the importance of nutrient release from
the nutrient pool in decomposing organic matter for ensuring uninter-
rupted mineral cycling in an ecosystem. The significance of nutrient release
is even more evident when considering not only the pools but also the
fluxes of nutrients in a forest. Figure 5 shows cycles of selected nutrients
in a few European forests. Note the relatively small nutrient retention in
plant biomass in comparison to nutrient uptake from soil and return with
litter fall.
The fragile balance between availability of nutrients for building new
organic matter and their return to the soil mineral pool can be relatively
easily lost as a consequence of anthropogenic disturbances, as has been men-
tioned. The prime example of this is found in intensively exploited forests,
which need to be fertilized because large quantities of nutrients are with-
drawn with harvest of biomass. Similar problems of decrease in pools of
available mineral nutrients may also result from industrial pollution, which
frequently suppresses organic matter decay rate. More details on these
problems are found in Chapter 8.