V.N. Beklemishev

Vladimir Beklimishev (1890-1962) was an outstanding Russian zoologist and ecologist. His most famous book "Principles of comparative anatomy of invertebrates" has been translated into English and many other languages.

His ecological and theoretical papers were never translated into English and are unknown outside Russia. However, many of his ideas are not outdated. This page gives a short outline of 3 papers published by Beklemishev in 1951-1964. These papers (among others) were later re-printed in one book: "Biocoenological principles of comparative parasitology" published in Moscow in 1970 by Nauka.

Short Biography

Vladimir Beklemishev was born on October, 4, 1890, in Grodno (Russia). 1913 - graduated from Peterburg University
1920 - professor at Perm University
1932 - chair of the Department of Entomology at the Institute of Malaria and Medical Parasitology in Moscow
1934 - professor at Moscow State University
1945 - member of Academy of Medical Sciences of the USSR.
He died on September, 4,1962.

General principles of organization of life

Beklemishev, V.N. 1964. On general principals of organization of life. Bulletin of MOIP, section Biology, 69(2): 22-38.

An organism is characterized by form, not by matter, because matter is continuously renewed. Cuvier (1817) considered life as a morphoprocess. It is a form sustained in the metabolism flow.

Three types of changes:

  1. onthogenetic changes (growth, development)
  2. necrotic changes (decomposition of the entire body or its portions)
  3. propagational changes (reproduction)
Three types of morphoprocesses:
  1. cyclic (division of protozoa, bacteria)
  2. cyclic-terminal (metazoa have necrotic components in their life cycle)
  3. terminal (no reproduction per se, i.e. the biosphere).
Cycles are actually spirals at a closer look.

Life forms can be characterized by:

  1. Degree of organization = is the intensity of life processes and the quality of their regulation (these two are correlated). Living matter differs from non-living matter by the degree of organization (it is very low in non-living matter).
  2. Complexity of organization = number of components and interactions. Biosphere is very complex. However, systems of intermediate complexity (individuals) usually have the highest degree of organization.
  3. Non-living components. Living systems may have non-living components. Moreover, at the lowest levels all living systems consist of non-living (=poorly organized) components (water, organic and inorganic molecules). There is no "living matter"; instead there are different degrees of organization. Cloths, tools are parts of human organization, parts of the body.
  4. Integrity (independence) of organization. When less external support is needed for existence and function of organization, then the integrity increases. Integrity of organisms changes during their life-cycle. In some stages of morphoprocess, organisms become open (e.g. they need copulation for reproduction, or they need food, etc.). When integrity increases, then organizations become less dependent on their environment.
  5. Harmony of organization = degree of cooperativeness among the parts of the organization. For example, if existence of the whole is based on destruction of its parts, then the organization is not harmonious. Of course, Beklemishev’s examples are all from biology. If he tried to take examples from sociology then his life could be much shorter.

Classification of interactions among populations

Beklimishev, V.N. 1951. On classification of biocoenological (simphysiological) interactions. Bulletin MOIP, Section Biology. 56(5): 3-30.

Beklemishev considered 8 types of interactions among populations:

I. Direct topic interactions.

One species changes physical or chemical conditions of the environment for another species. This is a one-way interaction: the 1st species affects the 2nd one, but there is usually no effect of the 2nd species on the 1st one.


And many other examples...

II. Direct trophic interactions

A. Consumer eats the live body of another species.
a. Consumer eats the entire body
  1. predation (predator selects the prey)
  2. non-selected consumption (plankton eaters)
b. Consumer eats some portions of the body and usually does not kill it.
  1. parasitism, grazing of grass (consumed parts are not produced for consumption)
  2. eating fruits that were produced by a plant for consumption.
c. Consumer eats non-living portions of the body. Example: Mallophaga (insects) eat feathers on birds.
B. Consumer eats excretions of the producer.
a. Consumption goes on the body of the producer
  1. excretions are not produced for consumption (flies)
  2. excretions are produced for consumption (nectar in flowers)
b. Consumption does not go on the body
  1. excretions are discrete (caprophagous insects)
  2. excretions are dispersed in air or water.
C. Consumer eats the carcass of the producer
a. Consumption of selected large dead bodies
b. Consumption of non-selected small dead bodies (saprophagous organisms)

III. Direct phoresic interactions

This is transportation of one species by another. Examples: transportation of seeds on the body of mammals. In many cases phoresic interactions are associated with other interactions (topic or trophic).

IV Direct fabric interactions

This is when organisms of one species use for their construction activity bodies or excretions of carcasses of another species. Examples: ant nests are built of plant leaves or needles.

I-a Indirect topic interactions

Species-1 depends on environmental conditions.
Species-2 changes the environment for species-1.
Species-3 affects species 2 and in this way indirectly interacts with species-1.

Example: Pine seedlings die under the shade of fir threes. However, when bark beetles kill a bunch of firs then pines will start growing in this gap. Thus, bark beetles have an indirect topic effect on pine population.

II-a Indirect trophic interactions

A. Non-living food (2 components: consumer + modificator). Effect on food.
1. Change of access to food sources
2. Inter-specific competition for food
B. Living food (3 components: consumer - modificator - producer). Effect on producer.
3. Increasing access to food. Phytophagous fish eats algae in which mosquito larvae are hiding. As a result, mosquitoes become more intensively eaten by a predatous fish. Another example: deer dig the snow and improve access of birds to berries under snow.
4. Decrease access to food. Algae hide mosquitoes from their predators.
5. Increase in prey numbers. Forest defoliators increase the number of bark beetles and this will increase food for woodpeckers.
6. Decrease in prey numbers due to topic effects on prey.
7. Competition for live food. When 2 predator species feed on the same prey species, then predators have an indirect trophic interaction.
C. Living food (3 components: consumer - modificator - producer). Effect on consumer.
8. Protection from enemies (ants protect aphids from other predators), or increase in the number of enemies.
9. Destruction of enemies. Example: host plant - herbivorous insect - predators and parasites.
10. Multiple prey. One prey species may attract predators from another prey species.

III-a Indirect phoresic interactions.

These are changes in abundance of transporting species or changes in access to these species. Also includes competition for transportation.

IV-a Indirect fabric interactions

These are changes in abundance of the builder species or species that is used for construction. Also includes changes in access to these species and competition among species that build and those that are used as materials for construction.

Spatial and Functional Structure of Populations

Beklemishev, V.N. 1960. Spatial and Functional Structure of Populations. Bulletin MOIP, section Biology, 65(2): 41-50.

Population is a set of conspecific individuals which interact with each other and inhabit the same area which is more or less isolated from areas occupied by other populations of the species.

Types of species area:

  1. small and non-differentiated
  2. large and differentiated into numerous populations
  3. large but almost non-differentiated (a superpopulation)
Differentiation of population is usually a result of the heterogeneity of environment. Migration occurs in both differentiated and non-differentiated populations. However, in differentiated populations it is less intensive.

Populations can be classified into the following groups:

  1. Independent populations. Can persist without any immigration, and its numbers do not depend on immigration.
  2. Semi-dependent populations. Can persist without any immigration, however, its numbers depend on immigration. If there is no immigration, then the abundance is much lower than in the presence of immigration
  3. Dependent populations. Cannot persist without immigration, however, organisms can reproduce there. Reproduction is not sufficient to sustain population numbers.
  4. Pseudopopulation. Do not reproduce at all. All organisms are immigrants.
  5. Temporary populations. In contrast to the first 4 types, these populations always get extinct after a while. Re-colonization is a rare event and thus, for some period of time the population is absent in the area. Temporary populations may reproduce however it is not sufficient.
  6. Hemipopulations. These are populations that consist of individuals in specific stages. Only those species have hemipopulations that change their environment during the life-cycle. Example: a hemipopulation of dragonfly larvae in the lake.
Functional population complex (FPC) = a set of populations that exchange by migrants. FPC usually have one or several independent populations that capable to support a number of dependent or semi-dependent populations.

Comments: FPC is almost the same as a metapopulation. The only difference is that Beklemishev thought that at least one independent population is necessary for a FPC. Metapopulation theory assumes that all local populations can eventually become extinct. Beklemishev’s approach is closer to island biogeography models where there is at least one stable independent population on the mainland.

Superpopulations also have internal structure: there is a variation of population density over space. Patches with increased density he call "subpopulations". Some subpopulations are stable, some are not stable. The latter are supported by the immigration flow from stable subpopulations.

Population is a symmorphological and a symphysiological unit.

If you are interested in Beklemishev's works, please contact me and I will try to answer your questions.

Alexei Sharov 03/03/96