V.N. Beklemishev

• Short Biography
• General principles of organization of life
• Classification of interactions among populations
• Spatial and Functional Structure of Populations

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 However, many of his ideas are not outdated.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

General principles of organization of life

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)

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).

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

Beklemishev considered 8 types of interactions among populations:

I. Direct topic interactions.

Examples:

• Organisms of the 2nd species use organisms of the 1st species as a substratum. Example: lichen on the trunk of a pine.
• Organisms of the 2nd species use excretions of organisms of the 1st species as a substratum. This kind of direct topic interactions can be associated with the direct or indirect trophic interactions. Example 1: dung insects eat dung produced by mammals (combination of direct topic with direct trophic interaction). Example 2: many predatorous insects (Staphilinidae, Histeridae) eat other dung insects (combination of direct topic with indirect trophic interaction).
• Organisms of the 2nd species use carcasses of organisms of the 1st species as a substratum. Examples: insects in trunks of dead trees.
• Organisms of the 2nd species use products and constructions of organisms of the 1st species as environment. Example: insects in rodent holes.
• Water weeds in creeks decrease the current speed and this may be favorable or unfavorable for other species.
• Tree shade is favorable for some species of grasses and shrubs

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

IV Direct fabric interactions

I-a Indirect topic interactions

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.

• By non-living food Beklemishev meant inorganic chemicals which are utilized by plants, excretions and carcasses of organisms. When food is non-living, then it is not a population. Probably, this was the reason why only 2 components of the system are considered.
• Beklemishev distinguished between 2 kinds of indirect interactions: (1) effect of the modificator on the numbers of one of the interacting species, and (2) effect on the access of one interacting species to another. These 2 types were recently re-discovered by Wootton (1994, Ecology, 75: 1544-1551, and 1994, Ann. Rev. Ecol. Syst. 25: 443-466).

III-a Indirect phoresic interactions.

IV-a Indirect fabric interactions

Spatial and Functional Structure of Populations

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)

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.

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.

Alexei Sharov 03/03/96