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CONCLUSIONS

The development of a large complex of adaptations in phylogeny of honey bee is attended by evolution of sociality. It arose in the process of relationship strengthening of females with their progeny.

Family consolidation is accompanied by individual advantages and competitions between its membets. Selection was conducive to conservation and selective changeability and decrease of evolution plasticity. That is why reproduction of signs contriduting to improvement of family adaptation. It occurred the turning of bee family into elementary unit of natural selection.

In the process of development of sociality honey bee acquired specific form of families reproduction by means of their division. The separated part of a family develop new places. They may be located on a large distance from the former ones. So it is increased the area of living in the process of division of families followed by some change of living conditions. The process of developing in a new area is accompanied by appearance of morphophysiologic differentiations. The existence of territorial — mechanical isolation hastens the fixation of acquired signs and rise of new populations.

The basic functions, providing vital activity of families, are performed by worker bees. The number of worker bees in families is limited by the fertility of queens. It has zonal changeability expressed in increase of nember while moving to areas with cold climate. The increase of bees number in families provides the improvement of cold-hardening. But zonal tendency to changeability of bees number in families doesn't have general prevalence. Some populations that originared and live in zones of moderate and warm climate exist in families consisting of a large number of bees.

The development of the mechanizm of optimization' of queen's fertility is attended by relationship of two opposite directions of selection.

The selection for improvement of viability in different extreme situations is favourable to increase the number of bees in families. The increase of number of bees resulted in increase of trophic substrate consume by each family. It is the factir limiting size of families within the population area, resulting in reduction of range of population

in the process of formation of population the qeeen's fertility is established at the level, providing recovery of elimination of bees. Their number in families determined by the necessity of their long and stable function combines with the economic use of vital resources.

Combs are built by instinctively coordinated interaction of large groups of adult worker bees. They form the basis of nest construction providing the necessary conditions for development of family members and keeping food reserve. By regulation of cell construction of double-sided combs is ensured the economy of building materials and the rational use of the nest space. It is achieved the adequacy of conditions thar regulare ontogenetic transformation by hard stereotype of form, size and space orientation of cells. Cells for normal development of worker bees and drones differ mainly in size (volume). The developing queens need cells of rather large volume on a level with orientation Of longitudinal axes in direction of vector of gravitation. Worker bees and drones need horizontal orientation of cells at the stage of larva transformation into chrysalis.

The fact tnat bees live in shelters influenced their reaction on sun light and its component — ultra violet rays (UV). Tolerance for UV changes in ontogeny. Bees are characterized by minimal tolerance for UV at the stage of embryo and nave maximal tolerance for UV at the imaginal stage. Large exposure of adult bees results in reduction of length of their life because of intensification of the process of physiological getting old. It is possible that UV rays have some influence on season changeability in length of life of bees.

It is reduced in spring and summer period in the result of increased exposure, but in autumn and winter it increases with the reduction of exposure! The sun position is used by bees as a reference point for orientation in space. The beginning and finish of bee flies depend on change of day lighting.

Season reproductive activity of queens in cycle is attended by the dynamics of photoperiod. Adaptation of physiological processes to season dynamic of photoperiod provides the anticipatory reaction to typical changes of environmental^ situation. The before winter preparations of bees function on this basis.

Photoperiod regulations don't influence the restriction of area occupied by honey bees. Mostly it is restricted by thermal factor. The wide range of tolerance for this factor is connected with eurythermic of bee families. Mainly it is achieved by thermoregulation inside nest that makes it independent from unfavourable temperature changes outside.

Social and individual adaptations to thermal factor developed Recording to the principle of divergence. The reaction of a family or a group of bees on unfavourable change of temperature is expressed in means of ethological regulation for the most part. It is achieved the effective counteraction to cooling by reduction of heat loss. In the complex of individual reaction to temperature change it is dominating the change of metabolism as a result of it. Its hyperactivity stimulated by overheating reflects on intensification of water secretion, evaporation of which contributes to cooling. Hypotermia stimulates migration from zone with unfavourable temperature. Short but effective protection from cooling is provided by cold stupor. It is stimulated by low temperature that inhibit locomotion.

The energy resources are retained in the state of cold stupor owing to reduction metabolism to a minimum. Their expenditures lower with fall of temperature. The fall of temperature is limited by point of maximal supercooling when it is started the freezing of liquid fractions of bee's body leading to its death.

Ethological physiological protection mechanisms that are used by bees for overcoming long time unfavourable situations function on the basis of consumption of carbohydrate food. The length of life of bee families is limited by the quantity of this food when it can't be replenished. So the instincts of search, delivery and storage of food resources attained great protection in phylogeny of honey bee. Of great importance are the means of their economic use.

The development of zones with moderate and cold climate is attended by adaptation to hibernating when bees are in deep hypodynamics. It is caused by factors madifying physiological state of bee family. Low temperature is the most important environmental factor. Low temperature inhibits greatly the locomotion activity of bees while stimulating cold aggregation. Among inside factors it is mostly important the reproductive diapause. The adaptive meaning of reproductive diapause is connected with the fact that the necessity of thermostabilization inside nest space is removed with absence of brood. Water is one of factor regulating reproductive diapause. Bees reduce the consumption of water by the end of summer and beginning of autumn because of limited possibility to fly and decrease of their producing. This is applied by fall of water content in carbohydrate food in the result of its conversion into honey. The increasing defficiency of water inhibits hormonal function of a queen that results in indifferent attitude to worker bees. After finish of flies it occurs the accumulation of water generated in the process of metabolism in rectums during hibernation. In the result of resorption water comes into crops and dilutes the contained honey. Connected with this replenishment of water it is induced the renewal of reproductive activity of a queen. Water vapor saturation of inside nest space has the same effect.

It is attended the season dynamics of water content in carbohydrate food by changenbility of the length of bee life. The increase of water content in food induces the reduction of the life length.

The content of water vapor in bee hives depends on intensity of its excretions in the process of breathing, environmental temperature and air humidity. The inside nest C02 concentration also depends on temperature. It increases with fall of temperature because of dynamics of consolidation of bees in intercomb spaces and intensity of air exchange. The way of living in poorly ventilated shelters resulted in the development of good hypoxia tolerance. Active reaction of bees on C02 surplus stimulates the increase of the concentration to 1% in summer and 3% in winter. High C02 concentration intensifies in bees the process of physiological getting old. The gas mixture containing C02 1/3 of the volume has the anaesthetic effect. The anaesthetic effect intensifies the age changes and reduces the length of life.

The natural nest shelters (hollows of trees, mountain cracks etc.) have screening effect alongside with light, mechanical and some heat protection. This effect reduces tension of electric fields in bee home. The nature variables of electric fields achieve tension that irritates bees by guided current in periods of high thunder activity. So the approach of thunder front makes bees being outside to go back to their homes thus decreasing the possibility of elimination due to bad weather. At home the influence of electric fields falls that contributes to normalization of state of bees.

The use of different natural means of protection from unfavourable influence of environment, the development of mechanisms providing independence on it by accumulation of trophic reserves in nest and optimization of microclimate inside with the help of ethological-physiological means influenced the restrictions of environmental valency in embryos, larvae, chrysalis. Stabilization of inside nest conditions is attended by development of stenotropness. It is followed by stabilization for length and ontogenic transformation of embryonal and postembryonal stages of development resulting in not high variety of morphometric sings. Honey bee is distinguished for stability of the signs among other single living and social insects.

The environmental requirement of queens, worker bees and drones at the embryonal stage of development are the same. The optimal temperature ensuring minimal elimination is about 33—36° C. The lower and upper temperature limits characterized by mean lethality is 30° С and 38° С respectively. The same lethality effect gives the reduction of relative humidity to about 50% in optimal temperature.

On the stage of chrysalis the environmental requirements of queens, worker bees and drones have some differences. The optimal temperature for queens ranges between 33—34° C, for worker bees and drones — 33—35° C. The lower temperature limit that is characterized by mean lethal effectiveness for queens and drones is 29,5—30,0° С and for worker bees it is

  1. 29° C. The upper temperature limit that has the same eliminating effectiveness for worker bees and queens is
  1. 38,0° C, for drones — 36,0° C, respectively.

The length of development depends on temperature. Embryonal stage shortens 1,5 times under the influence of temperature rise 7—8° С from lower to upper limit of lital range. On the same way it is changen the stage of chrysalis. The temperature dependence of the rate of development is related with metabolism intensity that is expressed in activity of 02 consumption and C02 discharge.

The changeability of morphometric sings is determined by variety of environmental conditions and their ability to meet the needs of embryos, larvae and chrysalis. Irrespective of this departure of body size and its parts from the range of population changeability is possible only towards reduction. The possibility of underdeveloped organs increases with growth of eliminating effect of a factor that destabilizes ontogeny. Unfavoutable effect of many biotic and abiotic factors results in underdeveloped wings having no wing plates. Their size (when they are formed) is characterized by not good changeability. The size of structures that-form proboscis are characterized by maximal and external skeleton of abdomen by minimal changeability. It may not coinside the directions of changes of different body parts induced by adequate environmental factors. Temperature rise from lower to upper limit of vital range results in increase of the length of proboscis, the diameter of head and the size of mandibles. When developed in optimal temperature wing plates reach max size. Diameter of abdominal rings don't change as the width of tergites reduces with raise of temperature and of sternites respectively increases. Limited larvae results in the size reduction of their wings, tergites and length of proboscis. The same direction of changes is caused by U.H.F. — field and UV exposure.

At the imaginal stage of development effected by selected elimination for morphological signs correlated with viability and/or functional properties of organs it occurs the reduction of ranges of changeability. Among queens the individuals with abdomen of large size and high number of well-developed ovarioles have all probability to survive and leave progeny. Individuals with relatively small wings, abdomen and genitals are eliminated in the process of drones maturations. So individuals developed in optimal conditions can survive.

Selective elimination for signs, the possibility of rise of which grows with departure from optimal conditions, is greatly conducive to limiting of individual changeability. It becomes more hard the tendency of reduction of its range by selective reproduction of queens and drones that are developing in optimal conditions.

So selection while being conducive to improvement of ethological-physiological mechanisms of stabilization of inside nest homeostasis, intensifies the consolidation of the morphological similarity. Differencies occur mostly because of rise of interspecific differentiations in connection with adaptation to local conditions.

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Источник: Неизвестный. Экология МЕДОНОСНОЙ ПЧЕЛЫ 1990. 1990

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