Вестник Томского государственного университета. Биология. 2019; : 64-81
Использование морфо-анатомических признаков для выявления гибридных растений в зоне естественной гибридизации Populus laurifolia и P. nigra в Сибири, Россия
https://doi.org/10.17223/19988591/46/4Аннотация
Список литературы
1. Wang Z.S., Du S.H., Dayanandan S., Wang D.S., Zeng Y.F., Zhang J.G. Phylogeny reconstruction and hybrid analysis of Populus (Salicaceae) based on nucleotide sequences multiple single-copy nuclear genes and plastid fragments // PLoS ONE. 2014. Vol. 9. e103645. doi: org/10.1371/journal.pone.0103645
2. Du S.H., Wang Z.S., Ingvarsson P.K., Wang D.S., Wang J.H., Wu Z.Q. Multilocus analysis of nucleotide variation and speciation in three closely related Populus (Salicaceae) species // Molecular Ecology. 2015. № 24. PP. 4994–5005. doi: org/10.1111/mec.13368
3. Liu X., Wang Z., Shao W., Ye Z., Zhang J. Phylogenetic and taxonomic status analyses of the Abaso section from multiple nuclear genes and plastid fragments reveal new insights into the north America origin of Populus (Salicaceae) // Frontier Plant Science. 2017. e28101098. doi: org/10.3389/fpls.2016.02022
4. Suarez-Gonzalez A., Hefer C.A., Lexer C., Cronk Q.C.B., Douglas C.J. Scale and direction of adaptive introgression between black cottonwood (Populus trichocarpa) and balsam poplar (P. balsamifera) // Molecular ecology. 2018. Vol. 27(7). PP. 1667–1680. doi: org/10.1111/mec.14561
5. Vanden Broeck V.A., Villar M., Bockstaele V.E., Slycken V.J. Natural hybridization between cultivated poplars and their wild relatives: evidence and consequences for native poplar populations // Annals of Forest Science. 2005. Vol. 62. PP. 601–613. doi: 10.1051/ forest:2005072
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10. Климов А.В., Прошкин Б.В. Фенетический анализ Populus nigra, P. laurifolia и P. × jrtyschensis в зоне гибридизации // Вавиловский журнал генетики и селекции. 2018. № 22(4). С. 468–475. doi: 10.18699/VJ18.384
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17. Климов А.В., Прошкин Б.В. Популяционно-фенетическая структура тополя лавролистного Populus laurifolia Ledeb. в бассейне реки Томи // Сибирский лесной журнал. 2018. № 5. С. 62–75. doi: 10.15372/SJFS20180506
18. Šiler B., Skorić M., Mišić D., Kovačević B., Jelić M., Patenković A., Kurbalija Novičić Z. Variability of European Black Poplar (Populus nigra) in the Danube Basin. Novi Sad: Vojvodinašume. 2014. 128 p. Available at: http://www.danubeparks.org/files/1903_BlackPoplarMonography.pdf
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22. Lindtke D., Gompert Z., Lexer C., Buerkle C.A. Unexpected ancestry of Populus seedlings from a hybrid zone implies a large role for postzygotic selection in the maintenance of species // Molecular Ecology. 2014. № 23. PP. 4316–4330. doi: 10.1111/mec.12759
23. Christe C., Stölting K.N., Bresadola L., Fussi B., Heinze B., Wegmann D., Lexer C. Selection against recombinant hybrids maintains reproductive isolation in hybridizing Populus species despite F1 fertility and recurrent gene flow // Molecular Ecology. 2016. № 25(11). PP. 2482–2498. doi: 10.1111/mec.13587
24. Zeng Y.F., Zhang J.G., Duan A.G., Abuduhamiti B. Genetic structure of Populus hybrid zone along the Irtysh River provides insight into plastid-nuclear incompatibility // Scientific Reports. 2016. № 6. e27306416. doi: 10.1038/srep28043
Tomsk State University Journal of Biology. 2019; : 64-81
Using morphological and anatomical characteristics to identify hybrid plants in the area of Populus laurifolia and P. nigra natural hybridization in Siberia, Russia
https://doi.org/10.17223/19988591/46/4Abstract
In this research, we revealed the overlapping of the areas of P. laurifolia (Tacamahaca section) and P. nigra (Aigeiros section) in the Altai-Sayan mountain country. There is a process of natural crossing with the formation of a hybrid species P. × jrtyschensis in their common habitat. Hybrid zones of poplars are interesting for selection; therefore, careful study is required. There is no practical using of P. × jrtyschensis and they are not yet included in floristic lists and identification guides of Siberian regions. The last is related to the lack of clear morphological characteristics important for its identification. The aim of this study was to examine morphological and anatomical characteristics of P. nigra, P. laurifolia and P. × jrtyschensis to assess the possibilities of their using in natural hybrids identification.
We conducted the main part of the studies in the populations of P. nigra, P. laurifolia, and P. × jrtyschensis of the Tom River Basin. A part of the samples for anatomical studies was taken in the populations of the Biya and Katun river basins (See Table 1). The morphological characteristics of the vegetative organs were examined by the comparative morphological method. We studied such qualitative characteristics as surface shape of elongated coppice shoots, surface shape of elongated canopy shoots, types of shortened canopy shoots, morphology of leaf blades and petioles. We collected herbarium material from each sample plot, from 30 reproductively mature trees distant from each other, at the southern side of the middle part of the canopy with the aim of researching the variability of morphological characteristics. 15 fully developed intact leaves from each individual from the middle part of shortened shoots were collected. The length of the leaf blade (L), the distance between the widest and basic parts (A) were measured for analyzing the shape of each leaf blade. Form determination was performed by calculating the A / L index. We used the following ranges: < 0.25 - triangular; 0.25- 0.35 - egg-shaped triangular; 0.35-0.45 - egg-shaped; 0.45-0.65 - elliptical; 0.65 > - reversed egg-shaped. Branches with generative buds were collected from 10 models of each taxon in the spring before the beginning of flowering (at the end of March) for studying the location of generative buds on shoots and flower morphology. We detected morphological features at a magnification of 16.3×. Inflorescences were taken from cutting branches placed in vessels with water (at a temperature of 20…24°C) and fixed in a mixture of 96% ethyl alcohol and water in a 1: 1 ratio. The study of the fruit morphology was performed on 30 fruitful catkins selected from trees during the fruiting period. The leaves for morpho-anatomical studies of the petiole were collected randomly from three individuals from populations in the Tom, Biya and Katun river basins. The selection was carried out from the middle part of the crown of reproductively mature trees, with shortened shoots. The petioles were separated from the leaf blades of shortened shoots and fixed in a mixture of 96% ethyl alcohol, glycerol and water at the ratio of 1: 1: 1. Cutting sections were made in the upper part of the stem (fan). The cutting samples were stained with 2% safranin aqueous solution. Anatomical features were studied at a magnification of 100×. The reliability of the results was achieved by studying the petioles of 9 leaves from each model.
Coppice shoots of P. nigra have acylindrical form along the entire length, rarely angulated in the upper part. Elongated coppice shoots of P. laurifolia are always ribbed. The majority of examined coppice shoots of P. jrtyschensis are mostly ribbed on top and cylindrical on bottom shoots. This feature is important for identifying hybrids at the early stages of the population development. Elongated canopy shoots, in contrast to coppice shoots, are less informative, because even P. laurifolia has cylindrical, angular and ribbed shoots. The same variation is typical of hybrids. The reliable hybrid identification between the species of Aigeiros and Tacamahaca sections according to the nature of the shortened canopy shoots differentiation is possible due to morphological methods application. Species of Tacamahaca section have special shortened shoots, discoblasts, species of Aigeiros section do not have them, and shortened shoots of the canopy are represented only by leptoblasts. It should be noted that discoblasts are always inherited by hybrids and are preserved even with return crosses (See Fig. 1). The studied species have four lamina forms: triangular, ovate-triangular, ovate and elliptical. At the endogenous level, several lamina forms can be observed at one tree, but one of them sharply prevails. P. nigra is characterized by triangular and ovatetriangular leaves, the last prevails in all populations (78.0-96.0%). P. laurifolia has three types of leaf blades: ovate-triangular, ovate and elliptical. Ovoid leaves are most common (50.0-70.0%) form for the studied populations. P. × jrtyschensis is characterized by the presence of all forms noted in the parent species. In populations of hybrids, ovate-triangular leaves (55.0-82.0%) are predominant, triangular and elliptical leaves are distinguished by an extremely low frequency of occurrence. Only one form of the leaf top and fan on shortened shoots in the middle part of the canopy for all the leaves of one tree is always expressed. The leaves of P. nigra in all studied populations have an exceptionally elongated pointed tip and a wedge-shaped fan. The shape of P. laurifolia leaf top is pointed. The intermediate characteristics are typical of P. × jrtyschensis leaf blades: the pointed elongated tip and the rounded wedge-shaped fan. Staminate and pistil late flowers of P. nigra, P. laurifolia and P. × jrtyschensis differ in the structure of the disc, flowers and pubescence (See Fig. 2-3). The ranges of the stamens number of the studied species do not overlap, and these characteristics can be considered as reliable in their identification. The fruit of P. laurifolia is a 2-4-fold capsule, usually a 3-fold capsule; 2-fold capsules are located mainly at its base, and 4-fold are in the upper part. Fruits of P. nigra are represented only by 2-fold capsules. 2-fold capsules predominate for P. × jrtyschensis, 3-fold capsules account for 13.0- 22.0%, concentrating mainly in the upper part of the catkins. The petioles of P. nigra and P. laurifolia differ in the shape of the cross section and, especially, in the contour of the adaxial side (See Table 2). The upper side of P. nigra is always round, that of P. laurifolia is heart-shaped due to a well-defined groove. The conductive system of P. nigra is linear, that of P. laurifolia is arciform, formed by different rings of closed collateral conductive beams. P. × jrtyschensis inherits the characteristics of the parent species, but is well identified by the cross section of the stem, the contour of the adaxial side, and the form of the conducting system (See Fig. 4). Due to the availability of anatomical studies of the petiole and a quite early development of shortened shoots in young individuals, the use of petiole anatomical characteristics is, in our opinion, the most reliable method for hybrid identifying. The analysis of the morphological and anatomical characteristics of species in the natural hybridization zone showed that the parent species are well identified by morphology and petiole anatomy. The considered characteristics of hybrids are predominantly intermediate, although asymmetry of P. laurifolia is traced by the differentiation of canopy shoots. There is a clear division of adult trees into three groups - two are represented by parental species and one by hybrids - in the studied hybrid zone according to the morpho-anatomical characteristics.
The paper contains 4 Figures, 2 Tables and 24 References.
References
1. Wang Z.S., Du S.H., Dayanandan S., Wang D.S., Zeng Y.F., Zhang J.G. Phylogeny reconstruction and hybrid analysis of Populus (Salicaceae) based on nucleotide sequences multiple single-copy nuclear genes and plastid fragments // PLoS ONE. 2014. Vol. 9. e103645. doi: org/10.1371/journal.pone.0103645
2. Du S.H., Wang Z.S., Ingvarsson P.K., Wang D.S., Wang J.H., Wu Z.Q. Multilocus analysis of nucleotide variation and speciation in three closely related Populus (Salicaceae) species // Molecular Ecology. 2015. № 24. PP. 4994–5005. doi: org/10.1111/mec.13368
3. Liu X., Wang Z., Shao W., Ye Z., Zhang J. Phylogenetic and taxonomic status analyses of the Abaso section from multiple nuclear genes and plastid fragments reveal new insights into the north America origin of Populus (Salicaceae) // Frontier Plant Science. 2017. e28101098. doi: org/10.3389/fpls.2016.02022
4. Suarez-Gonzalez A., Hefer C.A., Lexer C., Cronk Q.C.B., Douglas C.J. Scale and direction of adaptive introgression between black cottonwood (Populus trichocarpa) and balsam poplar (P. balsamifera) // Molecular ecology. 2018. Vol. 27(7). PP. 1667–1680. doi: org/10.1111/mec.14561
5. Vanden Broeck V.A., Villar M., Bockstaele V.E., Slycken V.J. Natural hybridization between cultivated poplars and their wild relatives: evidence and consequences for native poplar populations // Annals of Forest Science. 2005. Vol. 62. PP. 601–613. doi: 10.1051/ forest:2005072
6. Eckenwalder J.E. Populus // Flora of North America Editorial Committee. 2010. № 7. PP. 5–14.
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23. Christe C., Stölting K.N., Bresadola L., Fussi B., Heinze B., Wegmann D., Lexer C. Selection against recombinant hybrids maintains reproductive isolation in hybridizing Populus species despite F1 fertility and recurrent gene flow // Molecular Ecology. 2016. № 25(11). PP. 2482–2498. doi: 10.1111/mec.13587
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