VARIABILITY OF MOSS GRAVIREACTIONS UNDER PHOSPHORUS DEFICIENCY IN THE SUBSTRATE
Рубрика:
| Lobachevska, O, Kyyak, N |
| Space Sci. & Technol. 2026, 32 ;(4):83-95 |
| https://doi.org/10.15407/knit2026.04.083 |
| Мова публікації: Українська |
Анотація: The infl uence of gravity was analyzed, and the morphological and functional responses of three moss species with diff erent
gravisensitivity — Ceratodon purpureus (Hedw.) Brid., Physcomitrium patens (Hedw.) Mitt., Ptychostomum imbricatulum (Mhll. Hal.) Holyoak & N. Pedersen were determined depending on the phosphorus content (Pi) in the substrate. Phosphates (Pi) are important components in the initiation of gravitropism: low Pi content reduces the gravisensitivity of protonemata and growth reversion aft er clinorotation. At the same time, low Pi concentrations lead to morphological modifi cations, including stimulation of branching, an increase in moss turf density and size within the upper 0.2 cm layer of the substrate. It was determined that a 25 μM Pi concentration is the minimum test level used to substantiate the eff ect of gravistimulation on phosphate activity. Based on the variability of the values of gravitropic angles of the lateral protonemal branches depending on phosphate content, fi ve angle classes ranging from 0 to 90° were distinguished, with a dominant group of angles at 42...57°. A signifi cant eff ect of phosphates on the bending angle (gravisensitivity) of C. purpureus and P. imbricatulum was observed, whereas the eff ect on P. patens was insignifi cant. The eff ect of low phosphorus concentrations on the angle group 74...89° of P. patens branches, which grow almost parallel to the surface substrate, enhances plagiotropic growth. Plagiotropism in the turf structure of P. patens with perennial protonemata enables an increase in turf area within the upper substrate layer and expands access to nutrients from other locations. This indicates the species-specifi c features in growth correction of mosses under Pi defi ciency and the effi ciency of gravisensitivity
in determining branch angle variation. Morphological modifi cations and dynamic changes in the gravitropic sensitivity of mosses under the infl uence of Pi are accompanied by physiological responses as a necessary condition for growth activity. Under phosphorus defi ciency, intracellular phosphatase activity in protonemal turfs increased, serving as a general indicator of metabolic status, and the stimulatory eff ect of phytohormones on phosphate metabolism was enhanced. It was determined that species-specifi c features of gravisensitivity in C. purpureus and P. patens may act as mediators of physiological processes during phosphate defi ciency and altered gravity. The variation in phosphatase activity is attributed to the ontogenetic characteristics of gametophyte developmental stages and the ecological association of mosses with specifi c substrates. |
| Ключові слова: acid phosphatase, clinorotation, gravisensitivity, gravitropic angle, phosphates, phytohormones |
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Physiology, 133(4), 1677—1690. doi: 10.1104/pp.103.032169
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Péret B., Desnos T., Jost R., Kanno S., Berkowitz O., Nussaume L. (2014). Root architecture responses: in search of phosphate.
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Plaxton W. C., Tran H. T. (2011). Metabolic adaptations of phosphate-starved plants. Plant Physiology, 156(3), 1006—1015.
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Press M. C., Lee J. A. (1983). Acid phosphatase activity in Sphagnum species in relation to phosphate nutrition. New Phytologist,
93, 567—573. doi: 10.1111/j.1469-8137.1983.tb02707.x
Proctor M. C. F., Smith A. J. E. (1995). Ecological and systematic implications of branching patterns in bryophytes. Experimental
and Molecular Approaches to Plant Biosystematics. Eds. P. C. Hoch, A. G. Stephenson. St Louis, MO: Missouri Botanical
Garden, 87—110.
Puga M. I., Mateos I., Charukesi R., Wang Z., Franco-Zorrilla J. M., Lorenzo L., Irigoyen M. L., Masiero S., Bustos R., Rogriguez
J., Leyava A., Rubio V., Sommer H., Paz-Ares J. (2014). SPX1 is a phosphate-dependent inhibitor of phosphate starvation
response 1 in Arabidopsis. Proc. Nat. Acad. Sci. USA, 111(41), 14947—14952. doi: 10.1073/pnas.1404654111
Rico-Reséndiz F., Cervantes-Pérez S. A., Espinal-Centeno A., Dipp-Álvarez M., Oropeza-Aburto A., Hurtado-Bautista E.,
Cruz-Hernandez A., Bowman J. L. K., Arteaga-Vazquez M., Cruz-Ramírez A. (2020). Transcriptional and morpho-physiological
responses of Marchantia polymorpha upon phosphate starvation. Int. J. Molecular Sci., 21, 8354, 1—25. doi: 10.3390/
ijms21218354
Ripetskyj R. T., Kit N. A., Chaban Ch. I. (1999). Infl uence of gravity on the photomorphism of secondary moss protonemata.
Adv. Space Res., 23, 2005—2010. doi: 10.1016/S0273-1177(99)00350-6
http://www.bryoecol.mtu.edu/ (Last accessed: 3 March 2026).
Goffi net B., Buck W. R., Shaw A. J. (2009). Morphology, anatomy, and classifi cation of the Bryophyta. Bryophyte Biology. Eds
A. J. Shaw, B. Goffi net. Cambridge: Cambridge University Press, 55—139.
Hanba Y. T., Do T., Takemura K., Kitajima S., Vyacheslavova A., Takata M., Beier M., Yokoi M., Shinozava A., Maeda A., Yasui
Y., Sakaguchi N., Kameishi R., Watanabe R., Okugava S., Ozaki R., Hirai S., Kamachi H., Kume A., Karahara H., Sakata Y.,
Onoda Y., Fujita T. (2025). First contact with greater gravity: moss plants adapted via enhanced photosynthesis mediated by
AP2/ERF transcription factors. Sci. Adv., 11, eado8664, 1—13.
Huang G., Zhang D. (2020). Th e plasticity of root systems in response to external phosphate. Int. J. Molecular Sci., 21, 5955.
doi: 10.3390/ijms21175955
Huang Y., Li Y., Zhou X., Yin B., Tao Y., Zhang Y. (2023). Moss patch size as a factor profoundly infl uencing soil nutrient
characteristics and multifunctionality of temperate desert in Central Asia. Ecological Indicators, 155, 110975. doi: 10.1016/j.
ecolind.2023.110975
Karban R. (2021). Plant communication. Annu. Rev. Ecology, Evolution, and Systematics, 52(1), 1—24. doi: 10.1146/annurevecolsys-
010421020045
Kern V. D., Sack F. D., White N. J., Anderson K., Wells W., Martin C. (1999). Spacefl ight hardware allowing unilateral irradiation
and chemical fi xation in petri dishes. Adv. Space Res., 24, 775—778.
Khorkavtsiv Y. D., Kordyum E. L., Lobachevska O. V., Kyyak N. Y., Kit N. A. (2015). Branching of Ceratodon purpureus protonemata
eff ected under altered gravity conditions. Ukrainian Botanical J., 72(6), 588—595.
Kume A., Kamachi H., Onoda Y. (2021). How plants grow under gravity conditions besides 1g: perspective from hypergravity
and space experiments that employ bryophytes as a model organism. Plant Molecular Biology, 107, 1—13. doi: 10.1007/
s11103-021-01146-8.
Lazarenko A. S., Vysotskaya E. I., Lesniak E. M. (1971). Chromosome Atlas of Leafy Mosses. Kyiv: Naukova Dumka.
Li J. W., Yin B. F., Suo F. Y., Zhou X. B., Tao Y., Zhang J., Li Y.G., Zhang Y. M. (2021). Eff ects of moss mortality on soil water
evaporation and infi ltration of moss-dominated biological soil crusts. Acta Ecologica Sinica, 41, 6533—6541.
Lin W. Y., Huang T. K., Leong S. J., Chiou T. (2014). Long-distance call from phosphate: systemic regulation of phosphate starvation
responses. J. Experimental Botany, 65(7), 1817—1827. doi: 10.1093/jxb/ert431
Liu Y., Wirén N. (2022). Integration of nutrient and water availabilities via auxin into the root developmental program. Current
Opinion in Plant Biology, 65, 102117. doi: 10.1016/j.pbi.2021.102117
Lobachevska O., Rabyk I., Kyyak N., Danylkiv I., Karpinets L., Sochanchak R., Beshley S., Shcherbachenko O., Baik O., Chomyn
I., Khorkavtsiv Ya., Khomyn L. (2022). Role of Bryophyte Cover in Revitalization of the Anthropogenic Transformed
Territories. Lviv: Levada.
Lobachevska O. V., Kyyak N. Ya., Khorkavtsiv Ya. D., Kit N. A. (2024). Gravisensitivity of Mosses: Th e Importance of Gravitational
Force in Ontogenesis. Lviv. URL: https://nvd-nanu.org.ua (Last accessed: 3 March 2026).
Lynch J. P., Brown K. M. (2001). Topsoil foraging: an architectural adaptation of plants to low phosphorus availability. Plant
and Soil, 237, 225—237. doi: 10.1023/A:1013324727040
Pearse A. G. E. (1960). Histochemistry Th eoretical and Applied. London.
Péret B., Desnos T., Jost R., Kanno S., Berkowitz O., Nussaume L. (2014). Root architecture responses: in search of phosphate.
Plant Physiology, 166, 1713—1723. doi: 10.1104/pp.114.244541
Plaxton W. C., Tran H. T. (2011). Metabolic adaptations of phosphate-starved plants. Plant Physiology, 156(3), 1006—1015.
doi: 10.1104/pp.111.175281
Press M. C., Lee J. A. (1983). Acid phosphatase activity in Sphagnum species in relation to phosphate nutrition. New Phytologist,
93, 567—573. doi: 10.1111/j.1469-8137.1983.tb02707.x
Proctor M. C. F., Smith A. J. E. (1995). Ecological and systematic implications of branching patterns in bryophytes. Experimental
and Molecular Approaches to Plant Biosystematics. Eds. P. C. Hoch, A. G. Stephenson. St Louis, MO: Missouri Botanical
Garden, 87—110.
Puga M. I., Mateos I., Charukesi R., Wang Z., Franco-Zorrilla J. M., Lorenzo L., Irigoyen M. L., Masiero S., Bustos R., Rogriguez
J., Leyava A., Rubio V., Sommer H., Paz-Ares J. (2014). SPX1 is a phosphate-dependent inhibitor of phosphate starvation
response 1 in Arabidopsis. Proc. Nat. Acad. Sci. USA, 111(41), 14947—14952. doi: 10.1073/pnas.1404654111
Rico-Reséndiz F., Cervantes-Pérez S. A., Espinal-Centeno A., Dipp-Álvarez M., Oropeza-Aburto A., Hurtado-Bautista E.,
Cruz-Hernandez A., Bowman J. L. K., Arteaga-Vazquez M., Cruz-Ramírez A. (2020). Transcriptional and morpho-physiological
responses of Marchantia polymorpha upon phosphate starvation. Int. J. Molecular Sci., 21, 8354, 1—25. doi: 10.3390/
ijms21218354
Ripetskyj R. T., Kit N. A., Chaban Ch. I. (1999). Infl uence of gravity on the photomorphism of secondary moss protonemata.
Adv. Space Res., 23, 2005—2010. doi: 10.1016/S0273-1177(99)00350-6
Smith A. J. E. (2004). Th e Moss Flora of Britain and Ireland. Cambridge: Cambridge University Press.
Toal T. W., Ron M., Gibson D., Kajala K., Splitt B., Johnson L. S., Miller N. D., Slovak R., Gaudinier A., Patel R., Lucas M., Provart
N. J., Spalding E. P., Busch W., Klienbenstein D. J, Brady S. M. (2018). Regulation of root angle and gravitropism. G3:
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Turner B. L., Baxter R., Ellwood N. T. W., Whitton B. A. (2001). Characterization of the phosphatase activity of mosses in relation
to their environment. Plant, Cell and Environment, 24, 1165—1176. doi: 10.1046/j.1365-3040.2001.00767.x
Turner B. L., Baxter R., Whitton B. A. (2003). Nitrogen and phosphorus fractions in soil solutions and drainage streams in
Upper Teesdale National Nature Reserve, northern England: implications of organic compounds for biological nutrient
limitation. Sci. Total Environment, 314—316, 153—170. doi: 10.1016/S0048-9697(03)00101-3
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Yang S.-Y., Lin W.-Y., Hsiao Y. M., Chiou T.-J. (2024). Milestones in understanding transport, sensing, and signaling of the
plant nutrient phosphorus. Th e Plant Cell, 36, 1504—1523. doi: 10.1093/plcell/koad326
Yang Z., Yuan Y. Guo J., Li J., Li J., Yu H., Zeng W., Huang Y., Vin L., Li F. (2024). Responses of soil C, N, P and enzyme activities
to biological soil crusts in China: a meta-analysis. Plants, 13(11), 1525. doi: 10.3390/plants13111525
Zeng H., Chen F., Zhu Q., Ali S., Du J., Zhu Y., Yi K. (2025). Th e interplay between phosphorus nutrition and abiotic stresses
in plants. J. Genetics and Genomics, 52, 1507—1523. doi: 10.1016/j.jgg.2025.08.008
Zhang Q., Yin B. F., Li J. W., Lu Y. X., Rong X. Y., Zhou X. B., Zhang B. C., Zhang Y. M. (2021). Eff ects of moss mortality on soil
enzyme activities in a temperate desert. Chinese J. Plant Ecology, 46, 350—361. doi: 10.17521/cjpe.2021.0127
Toal T. W., Ron M., Gibson D., Kajala K., Splitt B., Johnson L. S., Miller N. D., Slovak R., Gaudinier A., Patel R., Lucas M., Provart
N. J., Spalding E. P., Busch W., Klienbenstein D. J, Brady S. M. (2018). Regulation of root angle and gravitropism. G3:
Genes, Genomes, Genetics, 8, 49—58. doi: 10.1534/g3.118.200540
Turner B. L., Baxter R., Ellwood N. T. W., Whitton B. A. (2001). Characterization of the phosphatase activity of mosses in relation
to their environment. Plant, Cell and Environment, 24, 1165—1176. doi: 10.1046/j.1365-3040.2001.00767.x
Turner B. L., Baxter R., Whitton B. A. (2003). Nitrogen and phosphorus fractions in soil solutions and drainage streams in
Upper Teesdale National Nature Reserve, northern England: implications of organic compounds for biological nutrient
limitation. Sci. Total Environment, 314—316, 153—170. doi: 10.1016/S0048-9697(03)00101-3
Virchenko V. M., Nyporko S. O. (2022). Prodromus of Spore Plants of Ukraine: Bryophytes. Kyiv: Naukova Dumka.
Whitton B. A., Al-Shehri A. H., Ellwood N. T. W., Turner B. L. (2005). Ecological aspects of phosphatase activity in cyanobacteria,
eukaryotic algae and bryophytes. Organic Phosphorus in the Environment. Eds B. L. Turner, E. Frossard, D. S. Baldwin.
Wallingford: Commonwealth Agricultural Bureau, 205—241.
Yang S.-Y., Lin W.-Y., Hsiao Y. M., Chiou T.-J. (2024). Milestones in understanding transport, sensing, and signaling of the
plant nutrient phosphorus. Th e Plant Cell, 36, 1504—1523. doi: 10.1093/plcell/koad326
Yang Z., Yuan Y. Guo J., Li J., Li J., Yu H., Zeng W., Huang Y., Vin L., Li F. (2024). Responses of soil C, N, P and enzyme activities
to biological soil crusts in China: a meta-analysis. Plants, 13(11), 1525. doi: 10.3390/plants13111525
Zeng H., Chen F., Zhu Q., Ali S., Du J., Zhu Y., Yi K. (2025). Th e interplay between phosphorus nutrition and abiotic stresses
in plants. J. Genetics and Genomics, 52, 1507—1523. doi: 10.1016/j.jgg.2025.08.008
Zhang Q., Yin B. F., Li J. W., Lu Y. X., Rong X. Y., Zhou X. B., Zhang B. C., Zhang Y. M. (2021). Eff ects of moss mortality on soil
enzyme activities in a temperate desert. Chinese J. Plant Ecology, 46, 350—361. doi: 10.17521/cjpe.2021.0127