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«Ecology and diversity of forest ecosystems in the Asiatic part of Russia Proceedings from International conference, 14.2-18.2.2008, Kostelec nad ernmi lesy, Czech Republic, 2008 ...»

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Table No.1: Features of climatic parameters during the active and passive periods of vegetation for 2001-2003 (data of the arboretum meteostation of Mountain-Taiga station) Time of vegetation Parameters of klimate Active period Passive periol (april – june) (july – august) 2001 year Precipitations total, mm 142 Mean temperature, 0 С 12.2 25. Amount of the solar days 71 2002 year Precipitations total, mm 194 Mean temperature, 0 С 12.4 18. Amount of the solar days 61 2003 year Precipitations total, mm 77 Mean temperature, 0 С 12.8 18. Amount of the solar days 61 Average annual norm of 191 precipitations, мм In tab. 1 the vegetative period is conditionally divided by us on active (April - June), when all growth processes are maximal also passive (July - September), described stabilization, delay of growth and development of young runaways. It is executed owing to the big variability of weather in Primorski Krai. From data tab. 1 follows, that in the first active part of vegetation the smaller quantity of deposits, than in the second part of vegetation drops out considerably, and intensity of solar illumination in May remains at a high level. It does not promote activity top and lateral meristems, is brightly reflected at introductions in a delay of blooming of kidneys. All three vegetative periods something are characteristic. 2002 -"wet year", 2003 - droughty, 2001 - optimum, close to average values. The sums of precipitations a moisture 2001 and 2002 are close on values, at the same time 2001 is allocated on the maximal parameters of number of sunny days and a difference in average temperatures between Proceedings from International conference, 14.2-18.2.2008, Kostelec nad ernmi lesy, Czech Republic, active and passive the periods of vegetation. There is steady enough parameter on three years is an average temperature of air in the first phase of vegetation (12,2-12,80С). Usually cold and long spring, and also strong frost penetration of soil in a combination to spring deficiency of a soil moisture do not promote acceleration of physiological processes in the beginning of vegetation [20].

At discussion of the material of researches on formation of stomas surface in growing needles of species Pinaceae us were considered well-known situations: light exposure and temperature of air in a crone of trees accrue from below-upwards, and relative humidity of air, on the contrary decreases. Under V.R.Zalenskogo's law top leaves in comparison with bottom at one species of plants have difficulties in supply (transport) of water and the substances, aggravated in a drought. Other terminology has settled: top leaves - light, bottom - shadow. At the first green plastids are compact, but more finely, than at shadow leaves with friable, but large chlorophyll-bearing plastids. Light leaves (needles) have thicker a cuticle, at them also is well developed epicuticular wax in comparison with shadow needles [2,19,25]. Besides on epidermis leaves xeromorphous and light types it is usually formed more the stomas, than on shadow, necessary for intensive gas exchange [5].

In the form of diameters of needles of the investigated plants the characteristic distinctions are revealed. So, at photophilous picea koraiensis and picea pungens the figure of a cut of the needles, especially shined circle, comes nearer to a rhombus, and at a picea ajanensis is a shade-enduring plant this figure have the elipcoides forms. Applanate the needles is formed also by representatives of firs and related it Pseudotsuga menziesii, possessing rather greater shade tolerance, and the needles of a pinus densiflora, of pinus koraiensis and pinus strobus in a cut is similar to a triangle, but at the second the height of mesophyll prevails, and at last needles the most thin and narrow. The cross-section cut of a leaf of a pinus densiflora is approached to the concave semicircle.

The basic results of three-year researches under the quantitative analysis of stoma's formation on epidermis coniferous are shown in tab. 2.

Table No.2: Dynamics of formation of stomatal density on needles of the first year at 10 species of coniferous breeds in the Mountain Taiga's arboretum (2001-2003 yy.) Average stomatal density on 1 мм2 surfaces of needles in Time of various conditions gathering of the of illumination of a crone Plants material The shined top part of a crone The shaded bottom part of a crone 2001 2002 2003 2001 2002 Abies may 35,84 33,93 60,38 64,09 23,51 34, nephrolepis june 36,57 34,78 23,98 25,35 19,18 21, july 32,63 24,91 36,82 23,69 23,30 26, august 32,26 27,96 30,55 25,33 20,36 21, Average for 34,40 30,39 37,93 34,61 21,58 25, the period Abies may 115,80 38,36 54,76 124,37 31,70 45, hollophylla june 34,70 33,32 35,03 40,38 21,00 22, july 28,20 28,84 36,08 22,09 25,85 28, august 35,80 39,18 37,88 26,35 28,43 25, Average for 53,62 34,92 40,93 52,29 26,74 30, the period Abies may 106,21 40,41 34,50 78,23 56,03 37, concolor june 52,46 30,22 33,73 21,95 20,12 22, july 37,88 32,17 36,92 9,96 21,38 24, august 43,35 29,02 36,83 12,16 24,24 21, Average for 59,97 32,95 35,49 30,57 30,44 26, the period Pseudotsuga may 46,89 52,40 43,40 31,16 39,79 35, menziesii june 44,46 41,91 43,22 28,13 33,12 26, july 41,96 28,47 40,14 27,63 28,34 26, august 40,06 36,22 32,88 27,08 21,02 31, Average for 43,34 39,75 39,91 28,15 30,56 29, the period Pinus may 89,17 48,55 75,16 50,08 41,05 58, koraiensis june 76,90 55,60 78,16 59,66 49,18 62, july 72,17 53,82 68,34 54,13 55,98 60, august 66,14 65,78 91,40 62,84 59,82 55, Average for 76,09 55,93 78,13 56,67 51,50 59, the period Pinus may 58,32 52,65 ----- 49,34 37,75 ---- strobus june 61,52 56,60 43,26 45,37 38,25 36, july 41,91 42,52 48,47 41,68 29,09 25, august 49,30 44,26 69,31 61,64 49,82 53, Average for 52,76 49,00 53,68 49,50 51,63 38, the period Ecology and diversity of forest ecosystems in the Asiatic part of Russia Pinus may 31,16 95,05 ----- 56,77 ----- ---- densiflora june 127,70 105,01 107,27 81,48 61,29 53, july 143,07 89,10 137,23 101,02 49,29 49, august 146,00 79,60 100,82 83,71 65,31 47, Average for the period 119,98 92,17 115,10 80,74 58,63 49, Picea may 57,63 42,04 40,53 57,26 41,79 51, ajanensis june 54,67 44,16 51,73 45,22 42,86 40, july 37,13 43,69 52,94 48,10 47,07 35, august 39,08 42,47 49,66 35,86 41,35 46, Average for 47,12 43,09 48,71 46,61 43,26 43, the period Picea may 47,10 57,15 43,39 48,62 49,64 45, koraiensis june 32,20 46,53 32,99 37,53 39,97 30, july 46,96 35,97 42,86 40,54 37,71 35, august 38,45 44,31 35,41 36,12 41,93 33, Average for 41,17 45,99 38,66 40,75 42,31 36, the period Picea may 51,63 26,63 85,01 61,45 35,41 81, pungens june 40,17 27,03 41,61 45,70 34,58 20, july 32,41 28,80 45,70 33,34 35,43 35, august 41,48 49,02 38,78 32,06 33,61 35, Average for 41,42 32,87 52,77 43,13 34,75 43, the period The note: the crossed out section designates absence of the generated needles at the moment of gathering a material.

On the average for the period of supervision in most cases with increase in light exposure from below-upwards the density accrues, Before V.D.Tchernyshev [20] is shown for a picea ajanensis essential distinction between density stomas on needles in the top part of a crone (35 pieces) and in bottom, its less shined part (22 pieces). Obtained data tab. 2 testify and to deviations from the general law of increase density of stomas in epidermis of light needles. So, at a picea pungens, per 2001 2002 and pinus strobus 2002 (introduction plants) density of stomatal channels at light needles is a little bit less than pine, than at shadow. At a tree for picea ajanensis (a fragmentary specie) in 2002 light and shadow needles did not differ on this parameter. Also similar it is noted at a abies hollophylla 2001. The abies concolor т in 2002 forms needles of different illumination circles with an insignificant numerical difference. The given phenomenon is connected with active cyclonic activity (see tab. 1). The greatest deviations stomata's parameters of a P. pungens emphasize its expressed sensitivity to soil overwetting.

The stoma's thickness is quite logically maximal in young needles as stomatal the device is pawned in the middle of the last vegetative year which climatic conditions cause initial quantity of stomas apertures [10]. Therefore the majority of investigated species stomata's indexes has more than surfaces in the first, active part of the vegetative period - in May, June.

In August, in connection with the termination of the vigorous activity a meristems, there is a stabilization of values of quantity stomata's, characteristic for species of genus Abies, Picea, noted in the termination of growth of runaways. At the pine representatives having slowed down the morphogenesis of needles, the above-named law is shown not every year. Apparently, weather conditions of each year when individual light exposure of growing needles changed influenced.

Display maximal stomatal numerical sizes at the majority of species in more "droughty" part of the active vegetative period, though average temperatures of air at this time considerably below temperatures in second half of vegetation is interesting.

Concurrence of the minimal area of stomatas at the majority of the studied coniferous breeds with a maximum quantity of the dropped out deposits in 2002 Such communications is not less important demand the further researches, but they the tendency of increase a stoma's formations in connection with lack of a moisture (see tab. 2. do not contradict the leaf (needles) in conditions of sufficient humidifying has smaller density aperture known in anatomy and physiology to the thesis:, than the leaf generated in conditions of deficiency of a moisture. The same is possible to notice and for the most "droughty" 2003, when at the majority of species coniferous was observed.

If to compare stomata's indexes by a principle a local – introduction species between related plants of so essential difference us it is not revealed. Characteristic differences are noted between firs on years of vegetation: the P. concolor has more the stomas in 2001 of the maximal solar light. In it, possibly, it is shown more, than the ecological conditions, a hereditary attribute. There is a basis to believe, that the P. concolor on the anatomic-morphological structure is more photophilous, than shade-enduring a kind [21] with similar shaping crones in conditions shading, to characteristic pine representatives. Analyzing climatic features of its natural area, described contrasts of superfluous and insufficient soil humidifying at high humidity of air, it possesses greater amplitude of the adaptation to a level of humidifying of ground, but not air.

In the investigated three of pinuses and piceas the quantity of stomatas on epidermis is more than local species some, and time of blooming of kidneys for introductions objects is shifted in time for a little bit late terms [16]. It is clearly appreciable for the P. strobus and P. densiflora. For the soil moisture first availability, and to the second humidity of air are limiting has begun vegetations. On formation and accumulation of pigments green plastids the introduction trees also considerably lag behind local species [22]. It is explainable expenses of energy for introduction plants in the beginning of vegetation on adaptation of the photosynthetic device with greater size of processes of breath in the general stream of a metabolism.

Proceedings from International conference, 14.2-18.2.2008, Kostelec nad ernmi lesy, Czech Republic, Comparing with interpatrimonial differences, the increase in density stomatal cripts depending on insistence of plants to light is precisely traced. Epidermal stomas formations fir plants have the least quantity, and their maximal parameters are peculiar to pines.

Having studied quantitative characteristics the stomatal device, definition of the area stomatal poles, actually stomata's apertures (channels) and their cumulative area on 1 мм2 at the investigated plants in the middle of vegetation (tab. 3) has been made. If to compare with the sizes of the crypts areas by way of reduction on the first place will appear a pinus densiflora and a abies concolor, on the second - species - a P. hollophylla, a P. ajanensis, a P. nephrolepis, on the third too two species – P.

strobus and P. koraiensis and on the fourth place – Pseudotsuga menziesii, a picea ajanensis, a picea pungens. If to estimate these kinds in relation to a moisture the resulted number corresponds to increase of tolerance investigated coniferous to lack of a moisture of ground. The disorder of numerical values actually stomata's apertures is wider and not at all kinds is connected with the sizes of cripts. The smallest transpiration courses forms of epidermis umbraticolous A. nephrolepis and a photophilous species – P. koraiensis, and the greatest on the area of a P. densiflora. As a result, the most mesophytic species are appear two kinds - a pinus densiflora and a aabies concolor, and the steadiest to lack of a moisture - a picea pungens.

Having multiplied the area of stoma's apertures on quantity stomatas on мм2 epidermis surfaces a sheet plate, the apertures the average surface has been calculated on unit of the area of needles (tab. 3). Their numerical values are also correlations under the attitude of plants to abiotics factors of environment. In a number of experimental plants the minimal area of apertures has as much as possible shade-enduringa species – the A. neprolepis, a little bit greater - adapted to a dryness soils a P.

pungens and a P. koraiensis - a photophilous plant. Further the total areas of stomatal channels increase in process of increase of insistence of plants to light and a moisture with a maximum at a pinus densiflora. We shall notice also, that to the greatest area poles at this species there corresponds its fitness to lack of direct solar radiation, at a A. concolor - its the least shade tolerance in conditions of introduction.

Table No.3: Morphological characteristics of the stomatal device of coniferous plants Square of Square of Square's sum of Plants stomatal cript, stomatal channels, stomatal channels on mm2 10-5 mm2 10-5 mm2 10-5/mm Abies nephrolepis 103 16 Abies hollophylla 118 27 Abies concolor 141 46 Pseudotsuga menziesii 57 24 Picea ajanensis 115 29 Picea koraiensis 55 15 Picea pungens 55 20 Pinus koraiensis 72 24 Pinus strobus 87 36 Pinus densiflora 151 45 A.L.Fedorkov informs on a relative constancy of the sizes the stomatal device of pinus Sylvestris L. [15] also. However, unequivocal communication between the size of stomas, their density, and also under formation them a surface on epidermis can to not be. For reception of a full morphological picture for epiderma the account of a physiological role the epicuticular wax filling actually aperture channels of a pole which anatomic structure defines its water-keeping properties depending on humidity internal and an environment [18,19] is required.

Thus, dependence of parameters of development the stomatal device coniferous from the change of any factor of environment taken separately, is not always appreciable, as other ecological factor less essential in other conditions, can render appreciable influence now. For example, mass definitions of density stomata's formations at the picea ajanensis in top belt Sihote-Alin (mountain Oblachnaja) from 500 up to 1856 m above ур. The seas of statistical reliability have not found out [20].

In three items of selection the number of stomas on epiderma changed from 51 up to 57 pieces (a mistake of average values from 1.6 up to 2.7). It means, that, despite of obvious increase of solar radiation with height, decrease in temperature of environment as though levelled a stoma's density in epiderma P. ajanensis trees. At the same time we can ascertain: as a whole in crones of coniferous trees law of Zalenskii takes place.

CONCLUSIONS In the first, active part of vegetation the size of stoma's density at the majority of species coniferous is more, than in second, less active part of vegetation. On the shined (light) needles actually the density of stomatas is more, than on shaded (shadow).

The area apertures and their total surface on unit of a cover epidermis increases from need of plants for illumination and humidity of environment.

Researches reveal the tendency of increase a stomatal surfaces on needles of investigated plants in conditions of lack of a moisture and on the contrary.

Is competent to note also, that during adaptation to a climate of Primorski Krai the introdaction coniferous on a parameter stomatal density approach with local, despite of essential distinctions in terms of the beginning of the vegetative period at those and other species.

Apparently, density of stomatas on epidermal and even the sizes of stomatal poles can be applied as the markers estimating adaptation coniferous to various ecological factors, especially to modes of a moisture and solar radiation. But also these Ecology and diversity of forest ecosystems in the Asiatic part of Russia indexes are not quite sufficient for an estimation of an adaptive variety a stoma's complex if to not consider intensive formation of epicuticular wax on epidermis of coniferous species.

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