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ISSN 1995-4255, Contemporary Problems of Ecology, 2008, Vol. 1, No. 2, pp. 238–244. © Pleiades Publishing, Ltd., 2008.

Original Russian Text © S.A. Ostroumov, 2006, published in Sibirskii Ekologicheskii Zhurnal, 2006, Vol. 13, No. 1, pp. 73–82.

On the Concepts of Biochemical Ecology and Hydrobiology:

Ecological Chemomediators
S. A. Ostroumov
Lomonosov Moscow State University, Vorob’evy Gory, Moscow, 119992 Russia

Abstract—Earlier, the author published two books and some papers, in which he described conceptual
foundations of new scientific disciplines — biochemical ecology and biochemical hydrobiology. These trends
in research include studies of the role of chemical substances in interorganismal interactions, in communication
and regulation of supraorganismal systems. Another part of biochemical ecology concerns studies of the destiny
and transformation of external chemical substances when they interact with the organisms. Both natural and
man-made compounds are interesting for biochemical ecology. The basic concepts of biochemical ecology
include ecological chemomediators and ecological chemoregulators that have already been included in the body
of modern conceptions and are used in modern ecological literature. Application of biochemical ecology to
aquatic ecosystems creates the basis for development of biochemical hydrobiology.
DOI: 10.1134/S1995425508020100

The founding concepts of biochemical ecology were The importance of engaging chemical and biochem-
formulated in the book “Introduction to biochemical ical approaches for understanding ecosystems and bio-
ecology” [1], published in 1986. The author had offered cenoses has been underscored by many Vernadsky’s
a unified view of many facts on the border between followers working in ecology and hydrobiology.
ecology and biochemistry, based on a number of previ- The subsequent development of natural sciences
ous works [e.g., 2] and being in agreement with the ap- brought new questions from the border zone between
proach outlined in [3–5]. The further developments in ecology, chemistry, and biochemistry. Among them,
this field [e.g., 6–8] have endorsed the expediency of there is a prominent problem of how well integrated is
distinguishing this direction of research, rooted in Ver- living matter and the biosphere in general and what
nadsky’s doctrine of biosphere [9–12]. The present pa- are molecular mechanisms of regulation of biospheric
per summarizes several in-depth reviews [1, 8, 22] to equilibria (“ecological equilibria” in modern terms,
provide a brief analysis of this interdisciplinary field of which still need to be defined more precisely) and for-
science and introduce some key concepts and terms. mation of bodies of organic matter in the biosphere.
The idea of the importance of chemical approaches Studies of the features of ecological equilibria in the
for studies of ecological and biospheric processes and biosphere and the mechanisms of their maintenance and
relationships between organisms and the environment, disturbance, including those involving various organic
including the hydrosphere, was thoroughly developed compounds, especially secondary metabolites are cur-
in the works by Vernadsky [9–12] who stated that the rently gaining much importance [13–20]. The main
ocean in general should be regarded, at its every point, reasons of growing interest to these problems are multi-
as an unbreakable link between dead inert matter and faceted.
ever-changing living matter, chemically restructuring
the inert environment [9]. Following Vernadsky, the First, the advances in ecosystem science have under-
heterogeneous living matter in the ocean, the marine standably led to the accumulation of a wealth of data re-
life as a whole, can be viewed as a special mechanism, garding the structure and functioning of ecosystems,
changing the sea chemistry entirely [11]. their dynamics and stability. This information lays the
foundation for a new stage in the development of ecol-
According to Vernadsky [11], the Earth’s biosphere, ogy, with an increasing attention given to the factors
in addition to living, biogenic and inert matter, includes that regulate the formation of ecosystems’ structure,
bioinert matter, produced at the same time both by liv- their dynamics and functioning. A deep analysis of
ing organisms and by inert processes and representing the concept of ecologic equilibrium, still insufficiently
dynamic equilibrium systems of both [9]. In the bio- clear, is urgently needed [7].
sphere, we encounter various forms of biospheric mat-
ter: dead inert low-activity matter, alive disperse matter, Second, improvements in the chemical and bio-
very active chemically and geologically, and bioinert chemical techniques used to study living organisms and
matter, a natural structure from alive and inert matter. their environment reveal new features of biochemical
The alive matter is a form of activated matter [9]. processes and chemical compounds, especially second-


ary metabolites, mediating and regulating many inter- lators of such ecological processes as flows of energy
actions between organisms [6, 8, 21]. and matter through ecosystems.
Third, the human impact on the biosphere has risen Biochemical ecology can be appropriately restricted
sharply, including dangerous chemical pollution. This to the compounds of the second group. Furthermore,
process is understandably worrying because of the there is no need to encumber biochemical ecology with
growing disturbance of ecological processes and eco- many problems of the impact of abiotic environment on
logical equilibria in many areas of the biosphere animals and plants [3], e.g., uptake of inorganic nutri-
[15–20]. ents by living organisms, etc.
Fourth, the intensifying and developing aquaculture What, then, represents the main subject of study of
and the general rise in the consumption of biological re- biochemical ecology? However incomplete and prelim-
sources pose a problem of engineering artificial ecosys- inary our answer may be, it is reasonable to define this
tems with sufficient stability and capacity for regu- mainly as ecological interactions between organisms
lation. and their high-order systems (populations and commu-
Understanding of the mechanisms of maintenance nities) mediated by chemical compounds, mostly those
and disturbance of ecological equilibria (or ecosystem that act exclusively or preferentially as information-
stability) is critically dependent on the rapidly growing transmitting messengers or as regulators of ecological
information from the borderlines between ecology, bio- processes [1, 8]. Some aspects of anthropogenic impact
chemistry, chemistry, and physiology. upon the biosphere, such as chemical pollution, may
also be considered a subject of biochemical ecology
Some important aspects of these problems are sub- [13–20].
ject of study in the adjacent fields, such as physiology, The main objects of biochemical ecology are bio-
behavioral science, zoology, toxicology, biochemistry, chemically interacting organisms, populations, and
biophysics, etc. Usually, however, such studies are communities, the substances that mediate and regulate
mainly concerned only with facts from the particular the wide spectrum of ecological interactions (including
field and neglect general ecological points. These nar- both trophic and nontrophic interactions), and bio-
rowly specialized approaches to the chemical aspects of chemical reactions utilizing these compounds. Impor-
ecological interactions between organisms are neces- tantly, biochemical ecology regards chemical com-
sary but not sufficient for appreciation of the mecha- pounds and biochemical reactions first of all as compo-
nisms of regulation and destabilization of ecosystems. nents of ecosystems and participants of ecological pro-
The vast number of chemically mediated interactions, cesses in the biosphere [1, 8]. This conceptually distin-
influences, dependences, and signals in biogeocenoses guishes biochemical ecology from biochemistry, where
form a complicated large-scale system. Only painstak- the same compounds may be studied as products of
ing interdisciplinary studies can help us to understand intracellular metabolism disregarding the ecological
these systems and learn how to manage them. perspective and context.
At this point, it is pertinent to discuss biochemical The set of methods of biochemical ecology includes
ecology in general, its subject, object, and methods, and a wide assortment of techniques from biochemistry,
the characteristic features distinguishing it from other bioorganic chemistry, chemistry of natural compounds,
scientific disciplines. and various bioassays for the substances under study or
The field of science at the interface between ecol- their mixtures. In addition to traditional toxicological
ogy, chemistry, and biochemistry is sometimes called approaches, these assays may address physiological or
chemical ecology. According to Barbier [3], living or- behavioral reactions to the substances and preparations
ganisms from both plant and animal kingdom influence under study. Carefully done sophisticated bioassays are
their environment through intersecting action of vari- no less important for biochemical ecology than highly
ous molecules. These interactions may occur between sensitive methods of structural analysis of organic mol-
animals, between plants, between plants and animals, ecules. Finally, the concluding phase of many studies in
and between animals and plants. The impact of abiotic this field is chemical synthesis of the discovered natural
environment on animals and plants should also be con- compound following determination of its structure and
sidered. The study of such interactions and their ecological role. Therefore, the inventory of biochemi-
chemical effectors is the subject of chemical ecology. cal ecology also includes methods of organic synthesis.
This definition, analyzed from the ecological point The compounds studied by biochemical ecology, as
of view, is notable for dividing the interactions between a rule, are present in the organisms or excreted by them
living organisms into two important groups. First, there in much lower amounts than those used mainly as en-
are interactions involving substances and molecules as ergy sources or building materials. Many of classes to
sources of energy or building blocks for the organisms which these compounds belong have long been studied
that consume them. The second type of interactions en- biochemically with respect to their structure and metab-
gages molecules that serve mainly or exclusively as olism, most (but not all) of them belonging to the group
messengers transmitting certain information or as regu- of secondary metabolites.



Thus, the methods of biochemical ecology include another [1]. They include sex and aggregation phero-
modern techniques of purification and structural analy- mones, food attractants, etc.
sis of natural compounds, and methods of discovery of Ecological chemoregulators are compounds that
ecological functions of organic compounds, many of regulate behavior, physiology, and development of
which were developed or perfected in the past decades other organisms [1]. This term embraces many phero-
[1, 8]. High sensitivity of modern analytical assays is mones of both aquatic and terrestrial organisms, and
essential for revealing the interactions between organ- plant compounds acting on herbivorous animals (in-
isms. cluding arthropods) to disturb their development and
A very important group of signal or regulatory com- reproduction (e.g., plant-derived compounds acting as
pounds are pheromones. Many aspects of their role in juvenile hormones, molting hormones, phytoestrogens,
information transmission, inter-individual communica- etc.). Another group of ecological chemoregulators are
tion, and regulation, have been thoroughly reviewed compounds made by plants (allelopathic agents) and in-
[1]. Since then, many new data have appeared regard- hibiting other plant species. They act as natural herbi-
ing the role of pheromones in the ecology of aquatic cides, used by plants to compete and leading to
and terrestrial organisms. We believe that the canonical regulation of population density and species composi-
definition of pheromones, given in the works of Karl- tion in plant communities.
son and L&& uscher (reviewed in [1, 8]), needs to be Ecological chemoeffectors is the most common
amended. Taking new data and concepts into account, term, denoting all substances, both natural and anthro-
we suggest the following working definition of phero- pogenic, influencing the ecology of living organisms to
mones: a certain degree [1, 21, 22].
Pheromones are individual compounds or their mix- The action of many ecological chemomediators and
tures (complexes, sets, or combinations), which are se- chemoeffectors is mediated through their activation by
creted by organisms into their environment (aerial, enzymatic biochemical reactions. An example is given
aquatic, or onto the organisms’ outer surface), and by a number of substances of plant origin that protect
which have a function in signaling, information trans- plants against pathogenic fungi or phytophages. On the
mission, or influence on the respondents usually of the other hand, biochemical reactions and evolution of the
same biological species; pheromones can cause a de- respective enzymes are extremely important for detoxi-
fined reaction (behavioral, physiological, or develop- fication of potentially harmful compounds produced by
mental) in the respondents; pheromones can act as the organism’s ecological partners; co-evolutional ad-
stimuli activating or inhibiting some reactions, behav- aptation of fungi or insects and the plants they eat
iors, or physiological processes in the respondents [22]. proceeds in this way.
The appropriateness of this particular definition is The mechanisms of detoxification and biodegra-
based on the accumulated body of information on dation of xenobiotics are becoming especially signifi-
pheromones, discussed in [22]. cant because of the large-scale chemical pollution of
modern ecosystems. Detoxification and biodegradation
Various aspects of pheromones and the related com- of the polluting agents usually employ the same bio-
pounds have been investigated by many scientists in chemical mechanisms that the cells use to neutralize
Russia. V.E. Sokolov initiated studies in this direction natural toxic substances or xenobiotics. Importantly,
at the Severtsov Institute of Ecology and Evolution. chemical pollution of the environment can disrupt
The list of authors having actively published in the field chemical communication between organisms, which
of pheromones includes A. S. Isaev, K. V. Lebedeva, involves ecological chemomediators, chemoeffectors,
V. A. Minyailo, Yu. B. Pyatnova, M. Barbier, S. N. No- and chemoregulators, as defined above. Therefore, the
vikov, E. P. Zinkevich, and many others. Yu. P. Kozlov problems of chemical pollution [13–17] can also be
and colleagues have studied pheromones of fishes from regarded as part of biochemical ecology.
Lake Baikal. A new direction, sensory ecology, is ac-
tively developed [21]. The study of pheromones is a To conclude this brief description of the basic con-
very important direction in the field of chemical com- cepts of biochemical ecology, it is worthy to underscore
munication of fishes [6] and other aquatic and terres- its relations with the field that is increasingly termed
trial organisms [1, 22, 23]. chemical ecology [3] or ecological biochemistry [4].
The above discussion shows that biochemical ecology
In addition to pheromones, other terms can be sug- is distinct from and narrower than chemical ecology, if
gested to facilitate the characterization of roles of the latter is defined, following M. Barbier, as chemistry
chemical compounds in signaling and regulation of in- applied to ecology. Ecological biochemistry, some-
teraction between organisms and higher-order sys- times placed at the interface between ecology, chemis-
tems [1]. try, and biochemistry, should encompass biochemical
The most general term ecological chemomediators mechanisms of adaptation of organisms to their envi-
describes natural chemical compounds acting as media- ronment, and purely biochemical aspects of metabo-
tors in interactions between organisms, transferring in- lism of ecologically important compounds and me-
formation during signaling from one organism to chanisms of detoxification of xenobiotics. Biochemical



ecology is closer to ecology, while ecological biochem- 3) Lowering the use of pesticides through introduc-
istry is closer to biochemistry. tion of alternative ways of population control [1, 8].
It must be reiterated that the above discourse is not Such approaches would play a significant role in de-
an attempt to give some set definitions but rather an in- creasing human ecological footprint and lowering the
vitation to discuss a modern, highly dynamic and not amount of pollutants introduced into the biosphere.
yet finally shaped field at the interface between many It has been pointed out that one aspect very impor-
traditional scientific disciplines and directions, not lim- tant for conservation is making consumption of all
ited to those already mentioned. This multitude of sci- biospheric resources more “green” [18–20]. This
ence disciplines includes biogeochemistry, toxicology means a significantly higher level of ecological compe-
and ecotoxicology, aquatic chemistry, studies of tence both when exploiting natural resources (including
“ecometabolism” and marine biochemistry (e.g., works marine and freshwater bioresources) and when con-
of Khailov [2]), public health aspects of hydrobiology structing any new ecosystem, either aquaculture or
(see [8]), biochemical pharmacology, biochemical sys- biogeocenosis designed to processing and treatment of
tematics, chemistry of secondary metabolites and natu- sewage and polluted water. Knowledge of biochemical
ral compounds, etc. It will take time to provide final ecology of the respective organisms and ecosystems al-
definitions and draw firm borders between these lows one to see an important aspect of their self-regula-
disciplines; to do so now is premature. tion and maintenance of the stability of these
populations and biogeocenoses. Moreover, biochemi-
What can biochemical ecology contribute to solving cal ecology provides a key for practical management
the problems of nature conservation and management, and fine-tuning of ecological objects and processes, be-
biotechnology, and aquaculture? A brief answer (see cause it reveals regulatory ecological functions of vari-
chapter 7 in [8] for a detailed discussion) requires con- ous substances intrinsic for a given population or
sideration of both theoretical and practical aspects. ecosystem, and these compounds can be made and in-
From the theoretical point of view, biochemical ecol- troduced to biogeocenoses at will, varying time and
ogy brings into view yet another side of the material ba- place of the intervention. Some of these problems have
sis of ecological equilibria in the biosphere. If one been reviewed in several chapters of [1, 8].
compares energy and material flows in the ecosystems
to the street traffic, the compounds discussed here will No discussion of problems of biochemical ecology
play the role of traffic lights or traffic police. Biochemi- is complete without mentioning biotransformation of
cal ecology only starts to disentangle this complicated xenobiotics (both natural and anthropogenic) in organ-
system of ecological chemoregulators, which make an isms and ecosystems. This topic is given a concise treat-
important contribution to homeostasis and homeo- ment in chapter 6 of the book [8]. A more detailed
kinesis (stability sensu lato) of ecosystems, including analysis of these problems is beyond the scope of this
aquatic ones. brief review.
At this point, it is prudent to bring up several terms
Theoretical foundations of biotechnology will likely of chemical and biochemical ecology often mentioned
benefit from the ideas of biochemical ecology that ex- in the literature.
plain the raison d’etre of secondary metabolites and
other biologically active substances, which are among 1. Allelochemical (from the Greek allhlwu, or
the most important objects of biotechnology despite the allhlo — reciprocal, (to) each other) is a substance
lack of conceptual knowledge about their functions. that has a certain ecological value (but not as an energy
Furthermore, the biochemical-ecological approach source) for organisms belonging to biological species
seems appropriate in the analysis of principles and other than the organism producing this substance. This
mechanisms of formation and functioning of producer does not exclude the possibility that allelochemicals
cells important in biotechnology. can ultimately become important for their producers. A
wide class of substance can be considered allelo-
From the practical point of view, biochemical ecol- chemicals (chapters 2–4 and section 5.2 in [8]). At the
ogy together with other scientific disciplines can con- same time, they do not include such important sub-
tribute to lowering the pollution of the biosphere. Some stances as pheromones [3, 4, 9, 21, 23] and some auto-
ways to achieve this include: inhibitory allelopathic agents (chapter 3 in [8]).
1) Increasing the self-purification capacity of natu- 2. Exometabolite is a substance excreted to the envi-
ral and anthropogenic ecosystems, including aquatic ronment and generally believed to have some ecologi-
ones [13–17]. Biotechnology and genetic engineering cal significance (not simply a waste product). The
can be promising here to construct and manage micro- etymology of this term is interesting; it is derived from
organisms with an enhanced ability to destroy pollut- the Greek root ballw with two Greek prefixes. The
ants [1, 8]. first prefix, ec (ez before vowels) means separation or
2) Development and introduction of compounds and origin, also identifying movement from the inside to the
materials with increased capability for destruction in outside. The second prefix, meta, denotes commonal-
the environment. ity, joint action, intermediate, sequence in space or



time, change, or movement. The term stems from the 8) marking the environments and orientating the or-
word ballw, a polysemantic verb meaning “to throw” ganisms in time and space.
(as well as to put something on, to expel, to vent, to These functions are discussed in more detail in [1, 8]
pour, to sprinkle and even to get pregnant). with many examples of particular substances.
Exometabolites are a very wide and important class of It should be underscored that there are several ap-
substances, especially important for understanding proaches to classification of substances used in interac-
aquatic ecosystems [1, 5, 8]. However, this concept is tions between organisms. One attempt at such classi-
hard or impossible to apply to many ecologically im- fication was undertaken by Whittaker and Feeny in
portant substances of terrestrial ecosystems, which can 1971 (discussed in [1]). However, it has a substantial
act without being excreted to the environment, such as a disadvantage because it is based on an essentially an-
number of substances important for ecobiochemical thropomorphic concept of usefulness or benefit that or-
interactions of plants with fungi and animals. ganisms gain from producing a certain chemical.
3. Semiochemical is a substance that can de defined The concepts of ecological chemoregulators and
[8] as a chemical involved in signal, information, or chemomediators proved useful for development and
other similar nontrophic interactions between organ- modern interpretation of basic ideas of ecology [7, 24].
isms. In practice, however, this term is used more nar- Biochemical ecology and biochemical hydro-
rowly and applied to pheromones and some other biology. Application of the approaches of biochemical
substances, including kairomones and allomones ecology to aquatic ecosystems unavoidably leads to the
(chapter 5 in [8]); many toxic substances of high eco- idea of biochemical hydrobiology. This is appropriately
logical importance are not adequately described by this validated by the fact that aquatic organisms produce a
term because of its meaning. The first and fundamental great variety of biologically active substances [1, 25,
part of this word is derived from Greek shmerou 26], many of which play ecologically important roles of
(shmhiou) — characteristics, symbol, sign, token, cue, pheromones, toxins, repellents, antifeedants, etc. A re-
signal, banner, seal, etc. As such, the term “semio- view of the existing data on the role of chemical signals
chemical” is used in the cases when a chemical acts as a in the ecology of aquatic organisms is given in [1, 8].
signal, mark, or carrier of some information. At the The role of algal exometabolites is analyzed in [5]. In-
same time, many chemicals, e.g., toxins and chemo- teresting facts about the role of chemicals in the infor-
sterilants (chapters 2, 4 and 5 in [8]) and allelo- mation flow in freshwater ecosystems are covered in
chemicals (chapter 3 in [8]) use not as signals but more [27, 28].
directly, simply killing or sterilizing other organisms or Some of the data forming the basis of biochemical
inhibiting their growth. Thus, the term semiochemical hydrobiology make important difference in comparison
is not universal either and not as widely encompassing with biochemical ecology of terrestrial organisms. At
as some authors tend to define it. least three groups of such facts can be emphasized.
These important definitions illustrate the point that First, excretion of fatty acids and other lipids with
ecobiochemical interactions are very diverse. They are properties of surface-active substances (SAS) into wa-
analyzed in [1, 8], where ecological functions of chemi- ter is of great importance. These natural SAS can be ac-
cals are used as a base to systematically explore a great cumulated on the water/air interface and form a surface
multitude of organisms and substances combined into a film. The chemical composition of this biogenic film
single ecobiochemical continuum typical of biosphere determines many of its properties, oxygen transfer
and especially of hydrosphere. through it, heat balance of water surface and other im-
The wide diversity of producers of ecological portant parameters and processes [29]. Thus, biochemi-
chemomediators and a great number of chemical struc- cal ecology and hydrobiology becomes intertwined
tural scaffolds of secondary metabolites should not with ecological biophysics of aquatic systems and
conceal the fact that the variety of the functions of these hydrophysical processes of the ecosystem scale [29].
compounds is limited. The following list of the most Second, the area of active research is the transport of
important functions is by no means exhaustive [1, 8]: essential nutrients, including w3 polyunsaturated fatty
1) protection from consumers acids (PUFA), along food chains. Some available data
2) attacking the organisms used as food suggest that, under certain conditions, the composition
3) restricting the competitors for common resources and amounts of PUFA can significantly influence the
functioning of populations and ecosystems. Thus, sev-
4) attracting other organisms eral papers in this issue considering the problems of for-
5) regulation of interactions within a population, mation and transfer of PUFA along side chains
group or kin (Sushchik and colleagues, Dubovskaya and colleagues)
6) supplying precursor chemicals (e.g., precursors are of great interest.
of hormones or pheromones) Third, natural bodies of water contain dissolved vi-
7) conditioning the environment, including aquatic tamins [30]. Water in lakes and ponds have been found
habitats to contain vitamin B12 (0.001–0.85 mg/l), thiamine



(0.001–12 mg/l), biotin (0.0001–0.1 mg/l), niacin (up to I thank many colleagues — V. D. Fedorov, A. S. Isa-
3.3 mg/l), pantothenic acid (up to 0.26 mg/l), etc. (re- ev, V. A. Stonik, A. A. Zhuchenko, A. V. Kaluev,
viewed in [30]). The presence of vitamins is due to their A. O. Kasumyan, A. V. Oleskin, and others — for help-
production by some hydrobionts. For example, produc- ful discussions. The help of O. S. Ostroumov is appreci-
tion of vitamins by some cyanobacteria and associated ated.
heterotrophic satellite bacteria has been shown [31].
The presence of vitamins in the water can possibly
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