Tarchevsky I. A. Signal systems of plant cells / otv. ed. A.N. Grechkin. M.: Nauka, 2002.294 p.

UDC 633.11 (581.14: 57.04)

SPECIFIC FEATURES OF PLANT DISTRIBUTION IN WHEAT AGROPOPULATION BY CLASS OF VARIATION OF ELEMENT PRODUCTIVITY KOLOS

A. A. Goryunov, M. V. Ivleva, S. A. Stepanov

Growing conditions have a significant effect on the distribution of plants in the agropopulation of durum wheat according to the classes of variation in the number of spikelets, the number of caryopses of a spike and their weight. Among the varieties of the Saratov selection in an extreme year in terms of agroclimatic conditions, it is characteristic different number plants: old varieties - small classes, new varieties - large variation classes. Favorable agroclimatic conditions increase the number of plants attributed to higher classes of variation in the elements of ear productivity.

Key words: variety, spikelet, caryopsis, wheat.

FEATURES DISTRIBUTION OF PLANTS IN WHEAT AGROPOPULATION ON CLASSES OF THE VARIATION OF ELEMENTS EFFICIENCY OF THE EAR

A. A. Goryunov, M. V. Ivleva, S. A. Stepanov

Vegetation conditions essentially affect distribution of plants in agropopulation of durum wheat on classes of a variation number of spikelets, quantities kernels an ear and their weight. Among cultivars of the Saratov selection in the conditions of extreme year on agroclimatic conditions it is characteristic various number of plants: to age-old cultivars - the small classes, to new cultivars - the big classes of a variation. Favorable agroclimatic conditions raise number of the plants carried to higher classes of a variation of elements of efficiency of an ear.

Key words: cultivar, spikelet, kernel, wheat.

In the morphogenesis of wheat, according to researchers (Morozova, 1983, 1986), several phases can be distinguished: 1) morphogenesis of the apical part of the meristem of the embryonic bud, leading to the formation of the rudimentary main shoot; 2) morphogenesis of phytomeric elements of the rudimentary main shoot into plant organs, which determines the habitus of the bush. The first phase (primary organogenesis - according to Rostovtseva, 1984) defines, as it were, the matrix of the plant. As established (Rostovtseva, 1978; Morozova, 1986; Stepanov, Mostovaya, 1990; Adams, 1982), the features of the passage of the primary processes of organogenesis are reflected in the subsequent structure formation.

According to researchers (Morozova, 1986, 1988), the formation of phytomeres of the vegetative zone of the embryonic main shoot is a species-specific process, while the deployment of phytomere elements of the embryonic main shoot into functioning plant organs is a variety-specific process. The formation of phytomeres in the shoot generative zone is more variety-specific (Morozova, 1994).

The most contrasting is the significance of primary morphogenetic processes, i.e. the establishment and formation of phytomeres of the vegetative and generative zones of the wheat shoot and their subsequent implementation in the appropriate agroclimatic conditions when analyzing the structure of the yield according to the variation curves of the elements of shoot productivity (Morozova, 1983, 1986; Stepanov, 2009). This is preceded by selective accounting of the distribution of plants in their agropopulation according to the classes of variation of individual productivity elements, in particular, the number of spikelets, the number of caryopses in an ear, and the weight of caryopses in an ear.

Material and method

The research was carried out in 2007-2009. The following varieties of spring durum wheat of the Saratov selection were selected as objects of study: Gordeiform 432, Melyanopus 26, Melyanopus 69, Saratovskaya 40, Saratovskaya 59, Saratovskaya zolotistaya, Lyudmila, Valentina, Nik, Elizavetinskaya, Zolotaya Volna, Annushka, Krassar. The main observations and counts were carried out in small-plot field experiments in the fields of the station selection crop rotation of the Research Institute of Agriculture of the South-East and the Botanical Garden of SSU, the experiments were repeated 3 times. To carry out a structural analysis of the productivity of wheat varieties, at the end of the growing season, 25 plants from each replication were taken, which were then combined into a group, and 25 plants were randomly selected from it for analysis. The number of spikelets, the number of grains in the spikelets, and the mass of one grain were taken into account. Based on the data obtained, the

the peculiarities of the distribution of plants in the agropopulation of durum wheat were divided in accordance with the method of Z.A. Morozova (1983) according to the classes of variation of the elements of ear productivity. Statistical processing of the research results was carried out using the Excel Windows 2007 software package.

Results and its discussion

As our studies have shown, in the growing season of 2007, the main number of the main shoots of wheat varieties of Saratov selection by the number of spikelets was in the 2nd and 3rd classes of variation. Only a small number of plants were assigned to the 1st class - 4% (Table 1).

Table 1. The number of wheat shoots of Saratov varieties by classes of variation in the number of spikelets,% (2007)

Variety Class of variation

1st 2nd 3rd 4th 5th

Gordeiform 432 0 92 8 0 0

Meljanopus 26 4 76 20 0 0

Meljanopus 69 4 64 32 0 0

Saratovskaya 40 7 93 0 0 0

Ancient 4 81 15 0 0

Saratovskaya 59 4 76 20 0 0

Saratov golden 0 16 80 4 0

Lyudmila 8 44 48 0 0

Valentina 0 16 76 8 0

Nick 14 14 72 0 0

Elizavetinskaya 0 24 72 4 0

Golden wave 8 16 52 24 0

Annushka 0 20 64 16 0

Crassar 0 20 48 32 0

New 4 27 59 10 0

When analyzing varieties by groups, it was found that ancient varieties are characterized by a greater number of plants of the 2nd class of variation (81%) and a smaller number of plants of the 3rd class of variation (15%). For the group of new varieties, it was revealed that a larger number of plants belong to the 3rd class of variation (59%), some of the plants of the 4th class of variation (10%). It was found that in some new varieties the number of plants of the 4th class of variation is more than 10% - Krassar (32%), Zolotaya Volna (24%), Annushka (16%), and in some varieties their number is less than 10% (Valentina,

Saratovskaya golden, Elizavetinskaya) or not observed at all - Saratovskaya 59, Lyudmila, Nick (see Table 1).

In the growing season of 2008, which was distinguished by a more favorable agroclimatic state, among the varieties of the Saratov selection, both old and new, a greater number of plants by the number of spikelets were attributed to the 3rd class of variation. Not a single plant, as in the previous year, was presented in the 5th grade of the variation. It is characteristic that, in contrast to the new varieties of durum wheat, a greater number of plants of the 2nd class of variation was noted in the old varieties - 41% (Table 2).

Table 2. The number of wheat shoots of Saratov breeding varieties by classes of variation in the number of spikelets,% (2008)

Variety Class of variation

1st 2nd 3rd 4th 5th

Gordeiform 432 12 20 60 8 0

Meljanopus 26 4 36 56 4 0

Meljanopus 69 4 48 48 0 0

Saratov 40 4 60 28 8 0

Ancient 6 41 48 5 0

Saratov 59 28 48 24 0 0

Saratov golden 0 28 64 8 0

Lyudmila 8 44 48 0 0

Valentina 4 28 64 4 0

Nick 4 28 68 0 0

Elizavetinskaya 8 36 52 4 0

Golden wave 4 12 68 16 0

Annushka 0 28 60 12 0

Crassar 8 28 32 32 0

New 7 32 52.5 8.5 0

Among the new varieties of durum wheat, varieties were distinguished for which, as in the previous year, the presence of a part of plants in the 4th class of variation in the number of spikelets was characteristic - Krassar (32%), Zolotaya Volna (16%), Annushka (12%) , Saratov golden (8%), Valentina (4%), Elizavetinskaya (4%), ie, the same trend was observed as in the previous year, 2007 (see Table 2).

During the growing season of 2009, most of the wheat plants of the Saratov selection according to the number of spikelets were assigned to the 4th and 3rd classes of variation: new varieties - 45 and 43%, respectively, old varieties - 30 and 51%, respectively. It is characteristic that some

eye varieties are characterized by the presence of a larger relative to the average value of the number of plants of the 4th class of variation - Annushka (76%), Valentina (64%), Nick (56%), Zolotaya Volna (52%), Saratovskaya 40 (48%). Some varieties have plants of the 5th class of variation - Zolotaya Volna (12%), Krassar (8%), Lyudmila (8%), Gordeiform 432 and Saratovskaya 40 - 4% (Table 3).

Table 3. The number of wheat shoots of Saratov varieties by classes of variation in the number of spikelets,% (2009)

Variety Class of variation

Gordeiform 432 4 12 52 28 4

Meljanopus 26 4 36 44 16 0

Meljanopus 69 0 8 64 28 0

Saratov 40 0 ​​4 44 48 4

Ancient 2 15 51 30 2

Saratov 59 0 28 48 24 0

Saratov golden 4 8 72 16 0

Lyudmila 0 4 56 32 8

Valentina 0 0 36 64 0

Nick 4 4 36 56 0

Elizavetinskaya 4 12 40 44 0

Golden wave 0 4 32 52 12

Annushka 0 0 24 76 0

Crassar 0 8 40 44 8

New 1 8 43 45 3

Thus, the studies carried out have shown that the vegetation conditions significantly affect the distribution of plants in the agro-population according to the classes of variation in the number of spikelets. Among the varieties of the Saratov selection in an extreme year in terms of agroclimatic conditions, a larger number of plants are characteristic: old varieties - 2nd class, new varieties - 3rd class, and some of them of 4th class of variation. Under favorable agroclimatic conditions, the number of plants attributed to higher classes of variation in the number of spikelets of a durum wheat spike increases.

In the growing season of 2007, the number of the main shoots of wheat of the Saratov varieties according to the number of caryopses on a spike was in the 1st and 2nd classes of variation. Only some of the plants of some varieties were assigned to the 3rd, 4th and 5th classes (Table 4).

Variety Class of variation

1st 2nd 3rd 4th 5th

Gordeiform 432 96 4 0 0 0

Meljanopus 26 96 4 0 0 0

Meljanopus 69 92 8 0 0 0

Saratov 40 93 7 0 0 0

Ancient 94 6 0 0 0

Saratovskaya 59 80 20 0 0 0

Saratov golden 20 48 32 0 0

Lyudmila 0 64 24 12 0

Valentina 48 36 16 0 0

Nick 28 62 10 0 0

Elizavetinskaya 48 48 4 0 0

Golden wave 12 32 48 4 4

Annushka 52 36 12 0 0

Crassar 88 8 4 0 0

New 42 39 17 1.5 0.5

When analyzing varieties by groups, it was found that the old varieties are characterized by a greater number of plants of the 1st class of variation (94%) and a very insignificant proportion of plants of the 2nd class of variation (6%). For the group of new varieties, it was revealed that a larger number of plants of certain varieties also belong to the 1st class of variation - Krassar (88%), Saratovskaya 59 (80%), Annushka (52%), Valentina (48%), Elizavetinskaya (48%) ), individual varieties - to the 2nd class of variation - Lyudmila (64%), Nick (62%), Saratovskaya golden (48%), Elizavetinskaya (48%) or to the 3rd class - Golden wave - 48% ( see Table 3). In two cultivars, plants of the 4th class of variation in the number of caryopses were noted - Lyudmila (12%) and Zolotaya Volna - 4% (see Table 4).

During the growing season of 2008, which, as noted earlier, was distinguished by more favorable agro-climatic conditions, among the varieties of the Saratov selection, both old-time and new, a greater number of plants by the number of spikelets was attributed to the 2nd and 3rd classes of variation ... However, among the ancient varieties, two varieties differed in a large relative to average values ​​of the number of plants of the 2nd class - Saratovskaya 40 and Melyanopus 69 - 72 and 48%, respectively. Among the new varieties, 3 varieties also differed in a large relative to average values ​​of the number of plants of the 2nd class - Saratovskaya 59 and Valentina (72%), Lyudmila - 64%.

In contrast to the previous year, among the varieties of the Saratov selection, the presence of a certain number of plants attributed to the 4th class of variation in the number of spike grains is characteristic. This is especially characteristic of the varieties Melyanopus 26, Elizavetinskaya, Lyudmila, Gordeiform 432, Melyanopus 69, Nik, Annushka (Table 5).

Table 5. The number of wheat shoots of varieties of Saratov selection by classes of variation in the number of spike grains,% (2008)

Variety Class of variation

1st 2nd 3rd 4th 5th

Gordeiform 432 0 28 56 8 8

Meljanopus 26 0 24 48 24 4

Meljanopus 69 4 48 40 8 0

Saratov 40 0 ​​72 24 4 0

Ancient 1 43 42 11 3

Saratovskaya 59 20 72 8 0 0

Saratov golden 4 36 56 4 0

Lyudmila 0 64 24 12 0

Valentina 0 72 28 0 0

Nick 0 32 60 8 0

Elizavetinskaya 0 48 32 20 0

Golden wave 12 32 48 4 4

Annushka 4 44 40 8 4

Crassar 4 40 52 4 0

New 5 49 39 6 1

During the growing season of 2009, the distribution of wheat plants of Saratov varieties by the number of spikelets was different depending on the group belonging - old or new varieties. In the group of ancient varieties, most of the plants were assigned to the 3rd and 4th classes of variation - 42.5% and 27%, respectively. In two varieties, Melyanopus 26 and Melyanopus 69, plants of the 5th class of variation in the number of caryopses were observed (Table 6).

Among the new varieties, most of the plants were assigned to the 3rd and 2nd classes - 50.5 and 24%, respectively (Table 6). It is characteristic that some varieties are characterized by the presence of a larger relative to the average value of the number of plants of the corresponding class: 2nd class of variation - Saratovskaya 59 (56%), Elizavetinskaya (32%), Krassar (32%), Gordeiform 32 (28%), Saratovskaya zolotistaya (28%); 3rd grade variations - Valentina (72%), Annushka (60%), Krassar (56%), Saratovskaya 40 (52%), Nick (52%), Elizavetinskaya (52%); 4th grade variation - Z-

lottery wave (36%), Annushka (32%), Saratov golden and Lyudmila (20%). It is noteworthy that, in contrast to previous years, under the conditions of 2009, some of the plants of half of the varieties were in the 5th class of variation in the number of spike grains - Lyudmila, Nick, Zolotaya Volna, Annushka, Melyanopus 26 and Melyanopus 69 (see Table 6) ...

Table 6. The number of wheat shoots of Saratov breeding varieties by classes of variation in the number of spike grains,% (2009)

Variety Class of variation

1st 2nd 3rd 4th 5th

Gordeiform 432 12 28 28 32 0

Meljanopus 26 8 22 46 20 4

Meljanopus 69 12 8 44 32 4

Saratovskaya 40 4 20 52 24 0

Ancient 9 19.5 42.5 27 2

Saratovskaya 59 12 56 24 8 0

Saratov golden 4 28 48 20 0

Lyudmila 0 12 52 20 16

Valentina 4 20 72 4 0

Nick 8 24 52 8 8

Elizavetinskaya 4 32 52 12 0

Golden wave 4 12 40 36 8

Annushka 4 0 60 32 4

Crassar 12 32 56 0 0

New 6 24 50.5 15.5 4

The studies have shown that the growing season conditions significantly affect the distribution of plants in the agropopulation according to the classes of variation in the number of caryopses in a spike. Among the varieties of the Saratov selection in an extreme year in terms of agroclimatic conditions, a larger number of plants are characteristic: old varieties - 1st class, new varieties -1-, 2- and 3-rd classes, and some of them of the 4th variation class. Under favorable agroclimatic conditions, the number of plants attributed to higher classes of variation in the number of durum wheat spike grains increases.

In the growing season of 2007, the number of the main shoots of wheat of the Saratov breeding varieties by the weight of the ear caryopsis was in the 1st and 2nd classes of variation (Table 7).

When analyzing varieties by groups, it was found that for some ancient varieties, the number of plants of the 1st class of variation was

100% - Gordeiform 432 and Melyanopus 26.93% - Saratovskaya 40. The old variety Melyanopus 69 was significantly different in this regard, which is characterized by a greater number of 2nd class plants - 80%. For the group of new varieties, it was revealed that some varieties are characterized by a larger number of plants of the corresponding class relative to the average value: 1st class - Zolotaya Volna (96%), Saratovskaya 59 (80%), Krassar (76%), Annushka (68%); 2nd grade - Nick (52%), Lyudmila (48%), Saratovskaya golden (44%), Valentina and Elizavetinskaya (40%); 3rd class variations - Lyudmila (28%), Saratovskaya golden (24%), Nick (14%), Valentina - 12%. It is noteworthy that in two cultivars, Lyudmila and Valentina, plants of the 5th class were observed with variations in the weight of caryopses - 12 and 4%, respectively (see Table 7).

Table 7. The number of wheat shoots of Saratov breeding varieties by classes of grain weight variation,% (2007)

Variety Class of variation

1st 2nd 3rd 4th 5th

Gordeiform 432 100 0 0 0 0

Meljanopus 26 100 0 0 0 0

Meljanopus 69 4 80 16 0 0

Saratov 40 93 7 0 0 0

Ancient 74 22 4 0 0

Saratovskaya 59 80 16 4 0 0

Saratov golden 32 44 24 0 0

Lyudmila 12 48 28 12 0

Valentina 44 40 12 4 0

Nick 28 52 14 6 0

Elizavetinskaya 56 40 4 0 0

Golden Wave 96 4 0 0 0

Annushka 68 32 0 0 0

Crassar 76 20 4 0 0

New 55 33 9.5 2.5 0

During the growing season of 2008, a different number of plants of the corresponding class of variation in the weight of spike grains was observed. Among the old varieties of Saratov selection, a greater number of plants for this productivity element corresponded to the 2nd class of variation - 48%, among the new varieties - to the 3rd and 2nd classes of variation - 38 and 36%, respectively. A certain number of plants of the corresponding varieties are distributed in the 4th and 5th classes of variation (Table 8).

Variety Class of variation

1st 2nd 3rd 4th 5th

Gordeiform 432 12 48 32 4 4

Meljanopus 26 0 32 44 12 12

Meljanopus 69 16 60 20 4 0

Saratovskaya 40 24 52 12 8 4

Ancient 13 48 27 7 5

Saratov 59 48 48 4 0 0

Saratov golden 4 24 64 4 4

Lyudmila 12 48 28 12 0

Valentina 4 36 56 0 4

Nickname 12 44 32 12 0

Elizavetinskaya 8 36 36 20 0

Golden wave 8 28 40 20 4

Annushka 8 36 36 16 4

Crassar 4 28 48 20 0

New 12 36 38 12 2

Some Saratov varieties were distinguished by a large, relative to average value, representation of plants of the corresponding class of variation in the weight of spike grains: 1st class - Saratovskaya 59 (48%), Saratovskaya 40 (24%), Melyanopus 69 (16%); 2nd class - Melyanopus 69 (60%), Saratovskaya 40 (52%), Saratovskaya 59 and Lyudmila (48%, respectively), Nick (44%); 3rd grade - Saratovskaya golden (64%), Valentina (56%), Krassar (48%), Melyanopus 26 (44%); 4th grade - Elizavetinskaya, Zolotaya Volna and Krassar (20%, respectively); Variations of the 5th class - Melyanopus 26 - 12% (see Table 8).

Under the conditions of the 2009 growing season, most of the wheat plants of the Saratov breeding varieties were attributed to the 3rd and 4th classes of variation according to the weight of spike grains. Moreover, the average values ​​of the classes of variation of the group of old varieties and the group of new varieties differed significantly. In particular, the old varieties were distinguished by a large representation of plants of the 3rd and 4th classes of variation - 41.5 and 29.5%, respectively, the new varieties were distinguished by the predominant presence in the agropopulation of plants of the 4th and 3rd classes of variation - 44 and 26%, respectively. ... Attention is drawn to a significant number of plants of the 5th class of variation in the weight of spike grains, which is especially characteristic of the varieties Krassar (32%), Valentina (24%), Zolotaya Volna (20%), Saratovskaya 40-16% (Table 9) ...

Variety Class of variation

1st 2nd 3rd 4th 5th

Gordeiform 432 4 16 48 32 0

Meljanopus 26 4 28 38 18 12

Meljanopus 69 0 8 48 40 4

Saratov 40 4 20 32 28 16

Ancient 3 18 41.5 29.5 8

Saratovskaya 59 14 36 38 8 4

Saratov golden 4 8 28 52 8

Lyudmila 0 0 12 80 8

Valentine 0 8 28 40 24

Nick 8 20 28 36 8

Elizavetinskaya 0 20 24 44 12

Golden wave 0 16 32 32 20

Annushka 4 8 32 56 0

Crassar 0 8 12 48 32

New 3 14 26 44 13

As well as in other years, some varieties were distinguished by a large (relative to the average value) representation of plants of the corresponding class of variation in the weight of spike grains: 1st class - Saratovskaya 59 (14%); 2nd class - Saratovskaya 59 (36%), Melyanopus 26 (28%), Saratovskaya 40, Nik and Elizavetinskaya (20%, respectively); 3rd class variations - Gordeiform 432 and Melanopus 69 (48%, respectively), Saratov 59 (38%), Zolotaya Volna and Annushka (32%, respectively); Variation class 4 - Lyudmila (80%), Annushka (56%), Saratovskaya golden (52%), Krassar (48%), Melanopus 69-40% (see Table 9).

Thus, the studies carried out have shown that the distribution of plants in the agropopulation according to the classes of variation in the weight of the ear caryopses is significantly influenced by the vegetation conditions. For most of the old varieties, under extreme growing conditions, the number of plants of the 1st class is 93-100%, while the new varieties are favorably distinguished by a significant representation of plants of the 2nd and 3rd classes. Under favorable growing conditions, the proportion of plants of a higher class of variation increases, but for new varieties the same tendency persists - a greater number of plants of higher classes of variation in the weight of caryopses in an ear as compared to old varieties.

Morozova Z.A. Morphogenetic analysis in wheat breeding. M.: Moscow State University, 1983.77 p.

Morozova Z. A. Main patterns of wheat morphogenesis and their importance for breeding. M.: Moscow State University, 1986.164 p.

Morozova ZA Morphogenetic aspect of the problem of wheat productivity // Morphogenesis and plant productivity. M.: MGU, 1994.S. 33-55.

Rostovtseva ZP Influence of the photoperiodic response of a plant on the function of the apical meristem in vegetative and generative organogenesis // Light and morphogenesis of plants. M., 1978.S. 85-113.

Rostovtseva ZP Growth and differentiation of plant organs. M.: Moscow State University 1984.152 p.

Stepanov S.A., Mostovaya L.A. Evaluation of the productivity of the variety by the primary organogenesis of the wheat shoot // Production process, its modeling and field control. Saratov: Publishing house Sarat. University, 1990.S. 151-155.

Stepanov, S.A., Morphogenetic features of the implementation of the production process in spring wheat, Izv. SSU Ser., Chemistry, biology, ecology. 2009.Vol. 9, issue 1. S. 50-54.

Adams M. Plant development and crop productivity // CRS Handbook Agr. Productivity. 1982. Vol.1. P. 151-183.

UDC 633.11: 581.19

Yu.V. Dashtoyan, S. A. Stepanov, M. Yu. Kasatkin

Saratov State University N. G. Chernyshevsky 410012, Saratov, st. Astrakhanskaya, 83 e-mail: [email protected]

The features in the content of pigments of various groups (chlorophylls a and b, carotenoids), as well as the ratio between them in wheat leaves belonging to different shoot phytomers, have been established. The minimum or maximum content of chlorophylls and carotenoids can be observed in various leaves, depending on the conditions of the growing season of plants.

Key words: phytomer, chlorophyll, carotenoid, leaf, wheat.

STRUCTURE AND THE MAINTENANCE OF PIGMENTS OF PHOTOSYNTHESIS IN THE PLATE OF LEAVES OF WHEAT

Y. V. Dashtojan, S. A. Stepanov, M. Y. Kasatkin

Features in the maintenance of pigments of various groups (chlorophyll a and chlorophyll b, carotenoids), as well as parities between them in the leaves of wheat

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Plant resistance to pathogens is determined, as was established by H. Flor in the 50s of the 20th century, by the interaction of a complementary pair of genes of the host plant and the pathogen, respectively, the resistance gene (R) and the avirulence gene (Avr). The specificity of their interaction suggests that the products of the expression of these genes are involved in the recognition of the pathogen by the plant, followed by the activation of signaling processes to trigger defense reactions.

Currently, 7 signaling systems are known: cycloadenylate, MAP-kinase (mitogen-activated protein-kinase), phosphatidic acid, calcium, lipoxygenase, NADPH-oxidase (superoxide synthase), NO-synthase.

In the first five signaling systems, G proteins act as an intermediary between the cytoplasmic part of the receptor and the first activated enzyme. These proteins are located on the inner side of the plasmalemma. Their molecules are composed of three subunits: a, b and g.

Cycloadenylate signaling system. The interaction of the stressor with the receptor on the plasmalemma leads to the activation of adenylate cyclase, which catalyzes the formation of cyclic adenosine monophosphate (cAMP) from ATP. cAMP activates ion channels, including the calcium signaling system, and cAMP-dependent protein kinases. These enzymes activate proteins that regulate the expression of protective genes, phosphorylating them.

MAP kinase signaling system. The activity of protein kinases is increased in plants exposed to stress (blue light, cold, drying, mechanical damage, salt stress), as well as those treated with ethylene, salicylic acid, or infected with the pathogen.

In plants, a protein kinase cascade functions as a signaling pathway. Binding of the elicitor to the plasmalemma receptor activates MAP kinases. It catalyzes the phosphorylation of the cytoplasmic kinase MAP kinase, which activates MAP kinase upon double phosphorylation of threonine and tyrosine residues. It passes into the nucleus, where it phosphorylates transcriptional regulator proteins.

Phosphatidic acid signaling system. In animal cells, G proteins under the influence of a stressor activate phospholipases C and D. Phospholipase C hydrolyzes phosphatidylinositol-4,5-bisphosphate to form diacylglycerol and inositol-1,4,5-triphosphate. The latter releases Ca2 + from the bound state. Increased content calcium ions leads to the activation of Ca2 + -dependent protein kinases. Diacylglycerol, after phosphorylation with a specific kinase, is converted into phosphatidic acid, which is a signaling substance in animal cells. Phospholipase D directly catalyzes the formation of phosphatidic acid from lipids (phosphatidylcholine, phosphatidylethanolamine) membranes.

In plants, stressors activate G proteins, phospholipases C and D in plants. Consequently, the initial stages of this signaling pathway are the same in animal and plant cells. It can be assumed that phosphatidic acid is also formed in plants, which can activate protein kinases with subsequent phosphorylation of proteins, including transcription regulation factors.

Calcium signaling system. Exposure to various factors (red light, salinity, drought, cold, heat shock, osmotic stress, abscisic acid, gibberellin and pathogens) leads to an increase in the content of calcium ions in the cytoplasm due to an increase in imports from the external environment and release from intracellular stores (endoplasmic reticulum and vacuoles)

An increase in the concentration of calcium ions in the cytoplasm leads to the activation of soluble and membrane-bound Ca2 + -dependent protein kinases. They are involved in the phosphorylation of protein factors regulating the expression of protective genes. However, it has been shown that Ca2 + is able to directly affect the human transcriptional repressor without using the protein phosphorylation cascade. Also, calcium ions activate phosphatases and phosphoinosite-specific phospholipase C. The regulating effect of calcium depends on its interaction with the intracellular calcium receptor - the protein calmodulin.

Lipoxygenase signaling system. The interaction of the elicitor with the receptor on the plasmalemma leads to the activation of membrane-bound phospholipase A2, which catalyzes the release of unsaturated fatty acids, including linoleic and linolenic, from plasmalemma phospholipids. These acids are substrates for lipoxygenase. Substrates for this enzyme can be not only free, but also unsaturated fatty acids included in triglycerides. The activity of lipoxygenases increases under the action of elicitors, infection of plants with viruses and fungi. The increase in the activity of lipoxygenases is due to the stimulation of the expression of genes encoding these enzymes.

Lipoxygenases catalyze the addition of molecular oxygen to one of the carbon atoms (9 or 13) of the cis, cis-pentadiene radical of fatty acids. The intermediate and final products of lipoxygenase metabolism of fatty acids have bactericidal, fungicidal properties and can activate protein kinases. Thus, volatile products (hexenals and nonenals) are toxic to microorganisms and fungi, 12-hydroxy-9Z-dodecenic acid stimulated protein phosphorylation in pea plants, phytodienic, jasmonic acids and methyl jasmonate, through activation of protein kinases, increase the level of expression of protective genes.

NADPH-oxidase signaling system. In many cases, infection with pathogens stimulated the production of reactive oxygen species and cell death. Reactive oxygen species are not only toxic to the pathogen and the infected cell of the host plant, but also participate in the signaling system. Thus, hydrogen peroxide activates transcription regulation factors and the expression of protective genes.

NO-synthase signaling system. In the macrophages of animals that kill bacteria, along with reactive oxygen species, nitric oxide acts, which enhances their antimicrobial effect. In animal tissues, L-arginine is converted into citrulline and NO under the action of NO synthase. The activity of this enzyme was also found in plants, and the tobacco mosaic virus induced an increase in its activity in resistant plants, but did not affect the activity of NO synthase in sensitive plants. NO interacts with oxygen superoxide to form a very toxic peroxynitrile. With an increased concentration of nitric oxide, guanylate cyclase is activated, which catalyzes the synthesis of cyclic guanosine monophosphate. It activates protein kinases directly or through the formation of cyclic ADP-ribose, which opens Ca2 + channels and thereby increases the concentration of calcium ions in the cytoplasm, which in turn leads to the activation of Ca2 + -dependent protein kinases.

Thus, a coordinated system of signaling pathways exists in plant cells that can act independently of each other or in concert. A feature of the signal system is the amplification of the signal during its transmission. The activation of the signaling system in response to the action of various stressors (including pathogens) leads to the activation of the expression of protective genes and an increase in plant resistance.

Induced mechanisms: a) increased respiration, b) the accumulation of substances that provide stability, c) the creation of additional protective mechanical barriers, d) the development of a hypersensitivity reaction.

The pathogen, overcoming surface barriers and entering the conducting system and plant cells, causes plant disease. The nature of the disease depends on the resistance of the plant. According to the degree of resistance, four categories of plants are distinguished: sensitive, tolerant, hypersensitive and extremely resistant (immune). Let us briefly characterize them using the example of the interaction of plants with viruses.

In susceptible plants, the virus is transported from the initially infected cells through the plant, multiplies well and causes a variety of disease symptoms. However, in sensitive plants, there are protective mechanisms that limit viral infection. This is evidenced, for example, by the resumption of reproduction of the tobacco mosaic virus in protoplasts isolated from infected leaves of tobacco plants, in which the growth of infectivity has ended. Dark green zones formed on young leaves of diseased susceptible plants are characterized by a high degree of resistance to viruses. The cells of these zones contain almost no viral particles in comparison with the neighboring cells of the light green tissue. A low level of accumulation of viruses in cells of dark green tissue is associated with the synthesis of antiviral substances. In tolerant plants, the virus spreads throughout the plant, but reproduces poorly and causes no symptoms. In hypersensitive plants, the initially infected and neighboring cells are necrotic, localizing the virus in necrosis. It is believed that in extremely resistant plants, the virus reproduces only in the initially infected cells, is not transported through the plant, and does not cause disease symptoms. However, the transport of viral antigen and subgenomic RNAs in these plants was shown, and when the infected plants were kept at a low temperature (10-15 ° C), necrosis was formed on the infected leaves.

The mechanisms of resistance of hypersensitive plants are best studied. The formation of local necrosis is a typical symptom of an oversensitive plant response to pathogen attack. They arise as a result of the death of a group of cells at the site of the introduction of the pathogen. The death of infected cells and the creation of a protective barrier around necrosis block the transport of the infectious agent through the plant, prevent access to the pathogen of nutrients, cause the elimination of the pathogen, lead to the formation of antipathogenic enzymes, metabolites and signaling substances that activate protective processes in neighboring and distant cells, and in ultimately contribute to the healing of the plant. Cell death occurs due to the inclusion of the genetic program of death and the formation of compounds and free radicals that are toxic to both the pathogen and the cell itself.

The necrotization of infected cells of hypersensitive plants, controlled by the genes of the pathogen and the host plant, is a special case of programmed cell death (PCD). PCD is essential for the normal development of the body. So, it occurs, for example, during the differentiation of tracheid elements during the formation of xylem vessels and the death of the cells of the root cap. These peripheral cells die even when the roots grow in water, that is, cell death is part of the plant's development and is not caused by the action of the soil. The similarity between PCD and cell death during a hypersensitive reaction is that these are two active processes, the content of calcium ions in the cytoplasm also increases in a necrotizing cell, membrane vesicles are formed, the activity of deoxyribonucleases increases, DNA breaks down into fragments with 3'OH ends, condensation occurs nucleus and cytoplasm.

In addition to the inclusion of PCD, necrotization of infected cells of hypersensitive plants occurs as a result of the release of phenols from the central vacuole and hydrolytic enzymes from lysosomes due to the disruption of the integrity of cell membranes and an increase in their permeability. The decrease in the integrity of cell membranes is due to lipid peroxidation. It can occur with the participation of enzymes and in a non-enzymatic way as a result of the action of reactive oxygen species and free organic radicals.

One of the characteristic properties of hypersensitive plants is acquired (induced) resistance to re-infection by the pathogen. The terms were proposed: systemic acquired resistance (SAR) and localized acquired resistance (LAR). LAR is said to be when cells acquire resistance in the area immediately adjacent to local necrosis (distance of about 2 mm). In this case, secondary necrosis is not formed at all. Acquired resistance is considered systemic if it develops in the cells of a diseased plant far from the site of the initial introduction of the pathogen. SAR is manifested in a decrease in the level of accumulation of viruses in cells, a decrease in the size of secondary necrosis, which indicates the inhibition of the short-range transport of the virus. It is not clear whether LAR and SAR differ from each other, or whether this is the same process occurring in cells located at different distances from the place of primary penetration of the virus into the plant.

The acquired resistance is usually nonspecific. Plant resistance to viruses was caused by bacterial and fungal infections and vice versa. Resistance can be induced not only by pathogens, but also by various substances.

The development of SAR is associated with the spread through the plant of substances formed in the initially infected leaves. It was suggested that the SAR inducer is salicylic acid, which is formed during the necrotization of initially infected cells.

When a disease occurs, substances accumulate in plants that increase their resistance to pathogens. An important role in the nonspecific resistance of plants is played by antibiotic substances - phytoncides, discovered by B. Tokin in the 20s of the 20th century. These include low molecular weight substances of various structures (aliphatic compounds, quinones, glycosides with phenols, alcohols) that can delay the development or kill microorganisms. Standing out when onions or garlic are wounded, volatile phytoncides protect the plant from pathogens already above the surface of the organs. Non-volatile phytoncides are localized in the integumentary tissues and are involved in the creation of the protective properties of the surface. Inside cells, they can accumulate in vacuoles. In case of damage, the amount of phytoncides increases sharply, which prevents possible infection of the wounded tissue.

Phenols are also referred to as antibiotic substances of plants. In case of damage and disease, polyphenol oxidase is activated in cells, which oxidizes phenols to highly toxic quinones. Phenolic compounds kill pathogens and host plant cells, inactivate pathogen exozymes and are necessary for lignin synthesis.

Among viral inhibitors, proteins, glycoproteins, polysaccharides, RNA, phenolic compounds were found. There are infection inhibitors that directly affect viral particles, making them non-infectious, or they block viral receptors. For example, inhibitors from beet, parsley and currant juice caused almost complete destruction of tobacco mosaic virus particles, and aloe juice caused linear aggregation of particles, which reduced the possibility of particles entering cells. Reproduction inhibitors alter cellular metabolism, thereby increasing cell resistance, or inhibit viral reproduction. Ribosome-inactivating proteins (RIPs) are involved in plant resistance to viruses.

In hypersensitive tobacco plants infected with the tobacco mosaic virus, proteins have been found, originally called b-proteins, and now referred to as proteins associated with pathogenesis (PR-proteins) or proteins associated with resistance. The common name "PR proteins" suggests that their synthesis is only induced by pathogens. However, these proteins are also formed in healthy plants during flowering and various stressful influences.

In 1999, based on the amino acid sequence, serological properties, enzyme and biological activity, a standardized nomenclature of PR proteins for all plants was created, consisting of 14 families (PR-1 - PR-14). Some PR proteins have protease, ribonuclease, 1,3-b-glucanase, chitinase activities or are protease inhibitors. Higher plants do not have chitin. It is likely that these proteins are involved in plant protection from fungi, since chitin and b-1,3-glucans are the main components of the cell walls of many fungi, and chitinase hydrolyzes the b-1,3-bonds of chitin. Chitinase can also act as lysozyme, hydrolyzing peptidoglucans of bacterial cell walls. However, b-1,3-glucanase can facilitate the transport of viral particles along the leaf. This is because b-1,3-glucanase destroys callose (b-1,3-glucan), which is deposited in the cell wall and plasmodesmata and blocks the transport of the virus.

The composition of PR proteins also includes low molecular weight (5 kDa) proteins - modifiers of cell membranes of fungi and bacteria: thionines, defensins and lipid-carrying proteins. Thionines are toxic in vitro to phytopathogenic fungi and bacteria. Their toxicity is due to the destructive effect on the membranes of pathogens. Defensins have strong anti-fungal properties, but do not work against bacteria. Defensins from plants of the families Brassicaceae and Saxifragaceae suppressed the growth of fungal hyphae by stretching, but promoted their branching. Defensins from plants of the families Asteraceae, Fabaceae, and Hippocastanaceae slowed down hyphal elongation, but did not affect their morphology.

When plants are infected with pathogens, the activity of the lytic compartment of cells of sensitive and hypersensitive plants increases. The lytic compartment of plant cells includes small vacuoles - derivatives of the endoplasmic reticulum and the Golgi apparatus, functioning as primary lysosomes of animals, that is, structures containing hydrolases in which there are no substrates for these enzymes. In addition to these vacuoles, the lytic compartment of plant cells includes the central vacuole and other vacuoles equivalent to secondary lysosomes of animal cells that contain hydrolases and their substrates, as well as plasmalemma and its derivatives, including paramural bodies, and extracellular hydrolases localized in the cell wall and in the space between the wall and the plasma membrane.

The action of elicitor drugs is due to the presence of special biologically active substances in their composition. According to modern concepts, signaling substances or elicitors are biologically active compounds of various natures that, in very low dosages, measured in mil-, micro-, and in some cases even nanograms, cause cascades of various plant responses at the genetic, biochemical and physiological levels. Their impact on phytopathogenic organisms is carried out by influencing the genetic apparatus of cells and changing the physiology of the plant itself, giving it greater vitality, resistance to various negative environmental factors.

The relationship of plants with the surrounding world, as highly organized elements of ecological systems, is carried out by perceiving physical and chemical signals coming from the outside and correcting all processes of their vital activity by influencing the genetic structures, immune and hormonal systems. The study of plant signaling systems is one of the most promising areas of modern cell and molecular biology. In recent decades, scientists have paid much attention to the study of signaling systems responsible for plant resistance to phytopathogens.

Biochemical processes occurring in plant cells are strictly coordinated by the integrity of the organism, which is complemented by their adequate responses to information flows associated with various influences of biogenic and technogenic factors. This coordination is carried out due to the work of signaling chains (systems) that are intertwined into the signaling networks of cells. Signaling molecules turn on most hormones, as a rule, do not penetrate into the cell, but interact with receptor molecules of the outer cell membranes. These molecules are integral membrane proteins, the polypeptide chain of which permeates the thickness of the membrane. Various molecules that initiate transmembrane signaling activate receptors at nano concentrations (10-9-10-7 M). The activated receptor transmits a signal to intracellular targets - proteins, enzymes. This modulates their catalytic activity or the conductivity of ion channels. In response to this, a specific cellular response is formed, which, as a rule, consists in a cascade of sequential biochemical reactions. In addition to protein mediators, relatively small messenger molecules, which functionally mediate between receptors and the cellular response, can also participate in signal transmission. An example of an intracellular messenger is salicylic acid, which is involved in the induction of stress and immune responses in plants. After switching off the signaling system, messengers are rapidly degraded or (in the case of Ca cations) are pumped out through ion channels. Thus, proteins form a kind of "molecular machine", which, on the one hand, perceives an external signal, and on the other hand, has an enzymatic or other activity modeled by this signal.

In multicellular plant organisms, signal transmission is carried out through the level of cell communication. Cells "speak" in the language of chemical signals, which allows homeostasis of the plant as an integral biological system. The genome and signaling systems of cells form a complex self-organizing system or a kind of "biocomputer". The genome is the hard carrier of information in it, and signaling systems play the role of a molecular processor that performs the functions of operational control. We currently have only the most general information on the principles of this extremely complex biological education. The molecular mechanisms of signaling systems remain largely unclear. Among the solution to many issues, there is a need to decipher the mechanisms that determine the temporary (transient) nature of the activation of certain signaling systems, and at the same time, a long-term memory of their activation, which manifests itself, in particular, in the acquisition of systemic prolonged immunity.

There is a two-way relationship between signaling systems and the genome: on the one hand, enzymes and proteins of signaling systems are encoded in the genome, on the other hand, signaling systems are controlled by the genome, expressing some genes and suppressing other genes. This mechanism includes the reception, transformation, multiplication and transmission of a signal to the promoter regions of genes, programming of gene expression, changes in the spectrum of synthesized proteins and the functional response of the cell, for example, the induction of immunity to phytopathogens.

Various organic ligand compounds and their complexes can act as signaling molecules or elicitors showing induction activity: amino acids, oligosaccharides, polyamines, phenols, carboxylic acids and esters of higher fatty acids (arachidonic, eicosapentaenoic, oleic, jasmonic, etc.), heterocyclic and organoelement compounds, including some pesticides, etc.

Secondary elicitors formed in plant cells under the action of biogenic and abiogenic stressors and included in the signaling networks of cells include phytohormones: ethylene, abscisic, jasmonic, salicylic acids, and

also the polypeptide systemin and some other compounds that cause the expression of protective genes, the synthesis of the corresponding proteins, the formation of phytoalexins (specific substances that have antimicrobial action and cause the death of pathogenic organisms and affected plant cells) and, ultimately, contribute to the formation of systemic resistance in plants to negative environmental factors.

Currently, seven signaling systems of cells are most studied: cycloadenylate, MAP-kinase (mitogen-activated protein-kinase), phosphatidic acid, calcium, lipoxygenase, NADPH-oxidase (superoxide synthase), NO-synthase. Scientists continue to discover new signaling systems and their biochemical participants.

Plants, in response to attack by pathogens, can use different pathways for the formation of systemic resistance, which are triggered by different signaling molecules. Each of the elicitors, acting on the vital activity of a plant cell along a certain signaling pathway, through the genetic apparatus, causes a wide range of reactions, both protective (immune) and hormonal, leading to a change in the properties of the plants themselves, which allows them to withstand a whole range of stress factors. At the same time, an inhibitory or synergistic interaction of various signaling pathways intertwining into signaling networks occurs in plants.

Induced resistance in manifestation is similar to genetically determined horizontal resistance, with the only difference that its nature is determined by phenotypic changes in the genome. Nevertheless, it possesses a certain stability and serves as an example of phenotypic immunocorrection of plant tissue, since as a result of treatment with substances of an elicitor effect, it is not the plant genome that changes, but only its functioning, associated with the level of activity of protective genes.

In a certain way, the effects arising from the treatment of plants with immunoinducers are related to gene modification, differing from it in the absence of quantitative and qualitative changes in the gene pool itself. With artificial induction of immune responses, only phenotypic manifestations are observed, characterized by changes in the activity of the expressed genes and the nature of their functioning. Nevertheless, the changes caused by the treatment with phytoactivators of plants have a certain degree of resistance, which manifests itself in the induction of prolonged systemic immunity, which is maintained for 2-3 months or more, as well as in the preservation of the acquired properties by plants for 1-2 subsequent reproductions.

The nature of the action of a certain elicitor and the achieved effects are in the closest dependence on the strength of the generated signal or the dosage used. These dependences, as a rule, have a sinusoidal rather than rectilinear character, which may serve as evidence of switching signaling pathways during their inhibitory or synergistic interactions. high severity of their adaptogenic action. On the contrary, treatment with these substances in large doses, as a rule, caused desensitization processes in plants, sharply reducing the immune status of plants and leading to an increase in the susceptibility of plants to diseases.

Presidium of the Russian Academy of Sciences
JUDGED
2002 Bach Prize
Academician Igor Anatolyevich TARCHEVSKY
for the cycle of works "Signaling systems of plant cells"

Academician I.A. TARCHEVSKY
(Kazan Institute of Biochemistry and Biophysics KSC RAS, Bach Institute of Biochemistry RAS)

PLANT CELL SIGNALING SYSTEMS

I.A.Tarchevsky has been studying the effect of abiotic and biotic stressors on plant metabolism for almost 40 years. Over the past 12 years, most attention has been paid to one of the most promising areas of modern biochemistry and plant physiology - the role of cell signaling systems in the formation of stress. On this problem, IA Tarchevsky published 3 monographs: "Catabolism and stress in plants", "Plant metabolism under stress", and "Signal systems of plant cells". In 30 articles, IA Tarchevsky and co-authors published the results of studies of adenylate cyclase, calcium, lipoxygenase and NADPH-oxidase signaling systems of plant cells. The NO-synthase signaling system is being investigated.

Analysis of the peculiarities of plant catabolism under stress made it possible to draw a conclusion about the signaling function of “shipwreck fragments” - oligomeric products of degradation of biopolymers and “fragments” of phospholipids. The assumption made in this work about the elicitor (signaling) properties of cutin degradation products was later confirmed by foreign authors.

There were published not only works of an experimental nature, but also reviews in which the results of studies of signaling systems of plant cells by domestic and foreign authors were summed up.

Studies of lipid metabolism, begun in the author's laboratory by A.N. Grechkin and then continued by him in an independent laboratory, made it possible to obtain results of a priority nature, which significantly expanded the understanding of the lipoxygenase signaling cascade. The study of the influence of the intermediate of the NADPH-oxidase system, salicylic acid, on protein synthesis led to the conclusion about the reason for the long-established biological activity of another compound, succinic acid. It turned out that the latter is a mimetic of salicylate and its treatment of plants “turns on” signaling systems, which leads to the synthesis of salicylate-induced protective proteins and an increase in resistance to pathogens.

It was found that various exogenous stress phytohormones - jasmonic, salicylic and abscisic acids induce the synthesis of both the same proteins (which indicates that these hormones “switch on” the same signaling pathways) and proteins specific for each of them ( which indicates the simultaneous "on" and differing signal cascades).
For the first time in the world literature, I.A. Tarchevsky analyzed the functioning of all known signaling systems of cells in plants and the possibilities of their mutual influence, which led to the idea of ​​the existence of not isolated signaling systems in cells, but of a signaling network consisting of interacting systems.

A functional classification of pathogen-induced proteins was proposed, and a review of the features of the synthesis of these proteins “switched on” by various signaling systems was made. Some of them are participants in plant signaling systems, and their intensive formation provides an increase in the perception, transformation and transmission of elicitor signals into the genetic apparatus, others limit the nutrition of pathogens, others catalyze the formation of phytoalexins, the fourth - the reactions of strengthening the cell walls of plants, and the fifth cause apoptosis of infected cells. The functioning of all these pathogen-induced proteins significantly limits the spread of infection throughout the plant. The sixth group of proteins can directly affect the structure and function of pathogens, stopping or suppressing their development. Some of these proteins cause degradation of the cell wall of fungi and bacteria, others disorganize the functioning of their cell membrane, changing its permeability to ions, and still others suppress the work of the protein-synthesizing machine, blocking protein synthesis on the ribosomes of fungi and bacteria or acting on viral RNA.

Finally, for the first time, the work on the design of pathogen-resistant transgenic plants was summed up, and this review work was based on the above-mentioned classification of pathogen-induced protective proteins.Particular attention is paid to the results of studies using transgenic plants of the peculiarities of the functioning of cell signaling systems.

Studies of signaling systems in plant cells are not only of great theoretical importance (since they form the basis of molecular mechanisms of stress), but also of great practical importance, since they allow the creation of effective antipathogenic drugs based on natural elicitors and intermediates of signaling systems.

Timiryazev, Kostychev and Sissakian lectures by I.A. Tarchevsky (the latter co-authored with A.N. Grechkin), as well as speeches at international conferences (in Hungary, England, France, Poland, Turkey, Israel, India, Germany, etc.).

For research of one of the signaling systems - lipoxygenase, I.A. Tarchevsky and Corresponding Member of the Russian Academy of Sciences A.N. Grechkin in 1999 were awarded the V.A.Engelhardt Prize of the Academy of Sciences of the Republic of Tatarstan.

In many publications of I.A. Tarchevsky, his colleagues - Corresponding Member of the Russian Academy of Sciences A.N. Grechkin, Doctors of Biological Sciences F.G. Karimova, N.N. Maksyutova, V.M. Chernov, O.A. Chernov and candidate of biological sciences V.G. Yakovleva.

In 2001, on the initiative of I.A. Tarchevsky and with his participation as the chairman of the Organizing Committee, the International Symposium on signaling systems of plant cells was held in Moscow.

LITERATURE

1. Tarchevsky I.A. Catabolism and stress in plants. The science. M. 1993.83 p.
2. Tarchevsky I.A. Plant metabolism under stress. Selected Works. Publishing house "Feng" (Science). Kazan. 2001.448 p.
3. Tarchevsky IA Signaling systems of plant cells. Moscow: Nauka, 2002.16.5 pp. (in the press).
4. Maksyutova N.N., Viktorova L.V., Tarchevsky I.A. The effect of ATP and c-AMP on the synthesis of proteins of wheat caryopses. // Physiol. biochem. cultures. plants. 1989. T. 21. No. 6. P.582-586.
5. Grechkin A.N., Gafarova T.E., Korolev O.S., Kuramshin R.A., Tarchevsky I.A. The monooxygenase pathway of linoleic acid oxidation in pea seedlings. / In: "Biological Role of Plant Lipids". Budapest: Akad. Kiado. New York, London. Plenum. 1989. P.83-85.
6. Tarchevsky I.A., Grechkin A.N. Perspectives of search for eicosаnoid analogs in plants. / In: "Biological Role of Plant Lipids". Budapest: Akad. Kiado. New York, London. Plenum. 1989. P.45-49.
7. Grechkin A.N., Kukhtina N.V., Kuramshin R.A., Safonova E.Yu., Efremov Yu.Ya., Tarchevsky I.A. Metabolization of coronary and vernolic acids in pea epicotyl homogenate. // Bioorgan. chemistry. 1990. Vol.16. No. 3.S. 413-418.
8. Grechkin A.N., Gafarova T.E., Tarchevsky I.A. Biosynthesis of 13-oxo-9 (Z), 11 (E) -tridecadienoic acid in pea leaf homogenate. / In: "Plant Lipid Biochemistry. Structure and Utilization ". London. Portland Press. 1990. P. 304-306.
9. Grechkin A.N., Kuramshin R.A., Tarchevsky I.A. Minor isomer of 12-oxo-10,15-phytodienoic acid and the mechanism of natural cyclopentenones formation. / In: "Plant Lipid Biochemistry. Structure and Utilization ". London. Portland Press. 1990. P. 301-303.
10. Tarchevsky I.A., Kuramshin R.A., Grechkin A.N. Conversation of α-linolenate into conjugated trienes and oxotrienes by potato tuber lipoxygenase. / In: "Plant Lipid Biochemistry. Structure and Utilization ". London. Portland Press. 1990. P. 298-300.
11. Grechkin A.N., Kuramshin R.A., Tarchevsky I.A. Formation of new α-ketol by hydroperoxide dehydrase from flax seeds. // Bioorgan. chemistry. 1991. T. 17. No. 7. S. 997-998.
12. Grechkin A.N., Kuramshin R.A, Safonova E.Y., Yefremov Y.J., Latypov S.K., Ilyasov A.V., Tarchevsky I.A. Double hydroperoxidation of linolenic acid by potato tuber lipoxygenase. // Biochim. Biophys. Acta. 1991. V. 1081. No. 1. P. 79-84.
13. Tarchevsky I.A. Regulatory role of biopolymer and lipid degradation. // Physiol. plants. 1992, vol. 39, no. 6, pp. 156-164.
14. Tarchevsky I.A., Maksyutova N.N., Yakovleva V.G. Effect of salicylic acid on protein synthesis in pea seedlings. // Plant Physiology. 1996. Vol.43. No. 5, pp. 667-670.
15. Tarchevsky I.A., Maksyutova N.N., Yakovleva V.G., Chernov V.M. Mycoplasma-induced and jasmonate-induced pea plant proteins. // Reports of RAS. 1996. Vol. 350, No. 4, pp. 544-545.
16. Chernov V.M., Chernova O.A., Tarchevsky I.A. Phenomenology of mycoplasma infections in plants. // Physiol. plants. 1996, vol. 43, no. 5. S. 721 - 728.
17. Tarchevsky I.A. On the probable reasons for the activating effect of succinic acid on plants. / In the book. " succinic acid in medicine, Food Industry, agriculture ". Pushchino. 1997.S. 217-219.
18. Grechkin A.N., Tarchevsky I.A. Lipoxygenase signaling system. // Physiol. plants. 1999. T. 46. No. 1. S. 132-142.
19. Karimova FG, Korchuganova EE, Tarchevsky IA, Abubakirova MR Na + / Ca + exchange in plant cells. // Reports of RAS. 1999. T. 366. No. 6. S. 843-845.
20. Karimova F.G., Tarchevsky I.A., Mursalimova N.U., Grechkin A.N. Effect of the product of lipoxygenase metabolism -12-hydroxydodecenoic acid on phosphorylation of plant proteins. // Physiol. plants. 1999. Vol. 46. # 1. S. 148-152.
21. Tarchevsky I.A. Interaction of signaling systems of plant cells, "switched on" by oligosaccharides and other elicitors. // "New perspectives in the study of chitin and chitosan." Materials of the Fifth Conference. M. Publishing house VNIRO. 1999.S. 105-107.
22. Tarchevsky I.A., Grechkin A.N., Karimova F.G., Korchuganova E.E., Maksyutova N.N., Mukhtarova L.Sh., Yakovleva V.G., Fazliev F.N., Yagusheva M.R., Palikh E., Khokhlova L.P. On the possibility of participation of cycloadenylate and lipoxygenase signaling systems in the adaptation of wheat plants to low temperatures... / In the book. “The edges of cooperation. On the 10th Anniversary of the Agreement on Cooperation between Kazan and Giessen Universities ”. Kazan: UNIPRESS, 1999.S. 299-309.
23. Tarchevsky I.A., Maksyutova N.N., Yakovleva V.G., Grechkin A.N. Succinic acid is a mimetic of salicylic acid. // Physiol. plants. 1999. T. 46. No. 1. S. 23-28.
24. Grechkin A.N., Tarchevsky I.A. Lipoxygenase signaling cascade of plants. // Scientific Tatarstan. 2000. No. 2. S. 28-31.
25. Grechkin A.N., Tarchevsky I.A. Cell signaling systems and genome. // Bioorganic chemistry. 2000. T. 26. No. 10. S. 779-781.
26. Tarchevsky I.A. Elicitor-induced signaling systems and their interactions. // Physiol. plants. 2000. Vol.47.No. 2.P.321-331.
27. Tarchevsky I.A., Chernov V.M. Molecular aspects of phytoimmunity. // Mycology and phytopathology. 2000. T. 34. No. 3. S. 1-10.
28. Karimova F., Kortchouganova E., Tarchevsky I., Lagoucheva M. The oppositely directed Ca + 2 and Na + transmembrane transport in algal cells. // Protoplasma. 2000. V. 213. P. 93-98.
29. Tarchevsky I.A., Karimova F.G., Grechkin A.N. and Moukhametchina N.M. Influence of (9Z) -12-hydroxy-9-dodecenoic acid and methyl jasmonate on plant protein phosphorylation. // Biochemical Society Transactions. 2000. V. 28. N. 6. P. 872-873.
30. Tarchevsky I.A. Pathogen-induced plant proteins. // Applied Microbiology and Biochemistry. 2001. T. 37. No. 5. S. 1-15.
31. Tarchevsky I.A., Maksyutova N.N., Yakovleva V.G. Influence of salicylate, jasmonate and ABA on protein synthesis. // Biochemistry. 2001. T. 66. N. 1. S. 87-91.
32. Yakovleva V.G., Tarchevsky I.A., Maksyutova N.N. Influence of NO donor nitroprusside on protein synthesis in pea seedlings. // Abstracts of International Symposium "Plant Under Environmental Stress". Moscow. Publishing House of Peoples' Friendship University of Russia. 2001. P. 318-319.
33. Yakovleva V.G., Maksyutova N.N., Tarchevsky I.A., Abdullaeva A.R. Influence of donor and inhibitor of NO-synthase on protein synthesis of pea seedlings. // Abstracts of International Symposium "Signaling systems of plant cells". Moscow, Russia, 2001, June, 5-7. ONTI, Pushchino. 2001. P. 59.