Abstract
Aims
Desert regions are regarded as highly sensitive to climatic changes. In arid and semi-arid desert, photosynthetic organisms from biological soil crusts are poikilohydric and sensitive to fluctuations in precipitation. How do physiological properties such as concentration of biochemical constituents and enzymes respond to a precipitation gradient from semi-arid to arid desert regions?
Methods
We sampled cyanobacteria and moss crusts from four desert regions with distinctly different amounts of annual rainfall. Subsequently, the contents of photosynthetic pigments, malondialdehyde (MDA), osmotic adjustment substances, and antioxidative enzyme activities were correlated with the means of annual precipitation, evaporation, and temperature at the various sites.
Results
Crust type, precipitation level, and their interaction had significant influences on many physiological properties (photosynthetic pigments, proline, soluble sugar, and superoxide dismutase). The contents of soluble protein, proline, and soluble sugar of cyanobacteria/moss crusts decreased with increasing precipitation level. Superoxide dismutase and catalase activities of cyanobacteria crusts decreased significantly with increasing annual precipitation. No significant variations in MDA were observed between different precipitation regions in the two crusts.
Conclusions
Among the environmental variables tested, the annual amount of precipitation had the strongest effect on the physiological properties of moss/cyanobacteria crusts in different regions. Crust type combined with particular precipitation level influenced the physiological properties of crusts. Moreover, both moss and cyanobacteria crusts exhibited strong physiological adaptability to changes in precipitation. This result needs to be incorporated into future ecological models, which will help in understanding the function and vulnerability of biocrusts in the face of climate change.
This is a preview of subscription content, log in via an institution to check access.
Similar content being viewed by others
Data availability
The datasets generated during and/or analyzed during the current study are available from the corresponding author on reasonable request.
Abbreviations
- BSCs:
-
Biological soil crusts
- Chl:
-
Chlorophyll
- Car:
-
Carotenoid
- MDA:
-
Malondialdehyde
- PVP:
-
Polyvinylpyrrolidone
- SOD:
-
Superoxide dismutase
- CAT:
-
Catalase
- SD:
-
Standard deviation
- ANOVA:
-
Analysis of variance
- MAP:
-
Mean annual precipitation
- MAT:
-
Mean annual temperature
- MAE:
-
Mean annual evaporation
References
Akcay UC, Ercan O, Kavas M, Yildiz L, Yilmaz C, Oktem HA, Yucel M (2010) Drought-induced oxidative damage and antioxidant responses in peanut (Arachis hypogaea L.) seedlings. Plant Growth Regul 61:21–28. https://doi.org/10.1007/s10725-010-9445-1
Almeselmani M, Deshmukh PS, Sairam RK, Kushwaha SR, Singh TP (2006) Protective role of antioxidant enzymes under high temperature stress. Plant Sci 171:382–388. https://doi.org/10.1016/j.plantsci.2006.04.009
Barker DH, Stark LR, Zimpfer JF, Mcletchie ND, Smith SD (2005) Evidence of drought-induced stress on biotic crust moss in the Mojave Desert. Plant Cell Environ 28:939–947. https://doi.org/10.1111/j.1365-3040.2005.01346.x
Beauchamp C, Fridovich I (1971) Superoxide dismutase: improved assays and an assay applicable to acrylamide gels. Anal Biochem 44:276–287. https://doi.org/10.1016/0003-2697(71)90370-8
Belnap J, Hawkes CV, Firestone MK (2003) Boundaries in miniature: two examples from soil. Bioscience 53:739–749. https://doi.org/10.1641/0006-3568(2003)053[0739:BIMTEF]2.0.CO;2
Bowker MA (2007) Biological soil crust rehabilitation in theory and practice: an underexploited opportunity. Restor Ecol 15:13–23. https://doi.org/10.1111/j.1526-100X.2006.00185.x
Bowker MA, Reed SC, Belnap J, Phillips SL (2002) Temporal variation in community composition, pigmentation, and Fv/Fm of desert cyanobacterial soil crusts. Microb Ecol 43:13–25. https://doi.org/10.1007/s00248-001-1013-9
Bradford MM (1976) A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding. Anal Biochem 72:248–254. https://doi.org/10.1006/abio.1976.9999
Bu CF, Wang C, Yang YS, Zhang L, Bowker MA (2017) Physiological responses of artificial moss biocrusts to dehydration-rehydration process and heat stress on the Loess Plateau, China. J Arid Land 9:419–431. https://doi.org/10.1007/s40333-017-0057-8
Büdel B, Darienko T, Deutschewith K, Dojani S, Friedl T, Mohr KI, Salisch M, Reisser W, Weber B (2009) Southern African biological soil crusts are ubiquitous and highly diverse in drylands, being restricted by rainfall frequency. Microb Ecol 57:229–247. https://doi.org/10.1007/s00248-008-9449-9
Cakmak I, Marschner H (1992) Magnesium deficiency and high light intensity enhance activities of superoxide dismutase, ascrobate peroxidase, and glutathione reductase in bean leaves. Plant Physiol 98:1222–1227. https://doi.org/10.1104/pp.98.4.1222
Celikkol Akcay U, Ercan Akca O, Kavas M, Yildiz L, Yilmaz C, Oktem HA, Yucel M (2010) Drought-induced oxidative damage and antioxidant responses in peanut (Arachis hypogaea L.) seedlings. Plant Growth Regul 61:21–28. https://doi.org/10.1007/s10725-010-9445-1
Chen XH, Duan ZH (2015) Impacts of soil crusts on soil physicochemical characteristics in different rainfall zones of the arid and semi-arid desert regions of northern China. Environ Earth Sciences 73:3335–3347. https://doi.org/10.1007/s12665-014-3622-x
Coe KK, Belnap J, Sparks JP (2012) Precipitation-driven carbon balance controls survivorship of desert biocrust mosses. Ecology 93:1626–1636. https://doi.org/10.1890/11-2247.1
Concostrina-Zubiri L, Martínez I, Rabasa SG, Escudero A (2014) The influence of environmental factors on biological soil crust: from a community perspective to a species level approach. J Veg Sci 25:503–513. https://doi.org/10.1111/jvs.12084
Dougill AJ, Thomas AD (2004) Kalahari sand soil: spatial heterogeneity, biological soil crusts and land degradation. Land Degrad Dev 15:232–242. https://doi.org/10.1002/ldr.611
Feng W, Zhang YQ, Jia X, Wu B, Zha TS, Qin SG, Wang B, Shao CX, Liu JB, Fa KY (2014) Impact of environmental factors and biological soil crust types on soil respiration in a desert ecosystem. PLoS ONE 9:e102954. https://doi.org/10.1371/journal.pone.0102954
Ferrenberg S, Reed SC, Belnap J (2015) Climate change and physical disturbance cause similar community shifts in biological soil crusts. PNAS 112:12116–12121. https://doi.org/10.1073/pnas.1509150112
Fischer T, Veste M, Wiehe W, Lange P (2010) Water repellency and pore clogging at early successional stages of microbiotic crusts in inland dunes, Brandenburg, NE Germany. CATENA 80:47–52. https://doi.org/10.1016/j.catena.2009.08.009
Garcia-Pichel F, Belnap J, Neuer S, Schanz F (2003) Estimates of global cyanobacterial biomass and its distribution. Algol Stud 148:213–227. https://doi.org/10.3103/S0095452709030049
Gong HJ, Zhu XY, Chen KM, Wang SM, Zhang CL (2005) Silicon alleviates oxidative damage of wheat plants in pots under drought. Plant Sci 169:313–321. https://doi.org/10.1016/j.plantsci.2005.02.023
Green TA, Sancho LG, Pintado A (2011) Ecophysiology of desiccation/rehydration cycles in mosses and lichens. Plant desiccation tolerance. Springer, Berlin, pp 89–120
Grote E, Belnap J, Housman D, Sparks J (2010) Carbon exchange in biological soil crust communities under differential temperatures and soil water contents: implications for global change. Global Change Biol 16:2763–2774. https://doi.org/10.1111/j.1365-2486.2010.02201.x
Guo Z, Ou W, Lu S, Zhong Q (2006) Differential responses of antioxidative system to chilling and drought in four rice cultivars differing in sensitivity. Plant Physiol Bioch 44:828–836. https://doi.org/10.1016/j.plaphy.2006.10.024
Guo YY, Zhang WH, He JF, Zhou JY, Yu HY (2012) Effects of water stress and seed mass on germination and antioxidative enzymes of Xanthoceras sorbifolia. Afr J Biotechnol 11:4187–4195. https://doi.org/10.5897/AJB11.3201
Hagemann M, Henneberg M, Felde VJ, Drahorad SL, Berkowicz SM, Felix-Henningsen P, Kaplan A (2015) Cyanobacterial diversity in biological soil crusts along a precipitation gradient, Northwest Negev Desert, Israel. Microb Ecol 70:219–230. https://doi.org/10.1007/s00248-014-0533-z
Hashempour A, Ghasemnezhad M, Ghazvini RF, Sohani MM (2014) Olive (Olea europaea L.) freezing tolerance related to antioxidant enzymes activity during cold acclimation and non acclimation. Acta Physiol Plant 36:3231–3241. https://doi.org/10.1007/s11738-014-1689-3
Heindel RC, Governali FC, Spickard AM, Virginia RA (2019) The role of biological soil crusts in nitrogen cycling and soil stabilization in Kangerlussuaq, west Greenland. Ecosystems 22:243–256. https://doi.org/10.1007/s10021-018-0267-8
Hu CX, Zhang DL, Huang ZB, Liu YD (2003) The vertical microdistribution of cyanobacteria and green algae within desert crusts and the development of the algal crusts. Plant Soil 257:97–111. https://doi.org/10.1023/A:1026253307432
Hui R, Li XR, Chen CY, Zhao X, Jia RL, Liu LC, Wei YP (2013) Responses of Photosynthetic properties and chloroplast ultrastructure of Bryum argenteum from a desert biological soil crusts to elevated ultraviolet-B radiation. Physiol Plantarum 147:489–501. https://doi.org/10.1111/j.1399-3054.2012.01679.x
Hui R, Zhao RM, Song G, Li YX, Zhao Y, Wang YL (2018) Effects of enhanced ultraviolet-B radiation, water deficit, and their combination on UV-absorbing compounds and osmotic adjustment substances in two different moss species. Environ Sci Pollut R 25:14953–14963. https://doi.org/10.1007/s11356-018-1689-8
Jia RL, Li XR, Liu LC, Gao YH, Li XJ (2008) Responses of biological soil crusts to sand burial in a revegetated area of the Tengger Desert, Northern China. Soil Biol Biochem 40:2827–2834. https://doi.org/10.1016/j.soilbio.2008.07.029
Kärkönen A, Kuchitsu K (2015) Reactive oxygen species in cell wall metabolism and development in plants. Phytochemistry 112:22–32. https://doi.org/10.1016/j.phytochem.2014.09.016
Kidron GJ, Vonshak A, Dor I, Barinova S, Abeliovich A (2010) Properties and spatial distribution of microbiotic crusts in the Negev Desert. Catena 82:92–101. https://doi.org/10.1016/j.catena.2010.05.006
Kidron GJ, Barinova S, Vonshak A (2012) The effects of heavy winter rains and rare summer rains on biological soil crusts in the Negev Desert. Catena 95:6–11. https://doi.org/10.1016/j.catena.2012.02.021
Kidron GJ, Monger HC, Vonshak A, Conrod W (2012) Contrasting effects of microbiotic crusts on runoff in desert surfaces. Geomorphology 139–140:484–494. https://doi.org/10.1016/j.geomorph.2011.11.013
Lan SB, Wu L, Zhang DL, Hu CX, Liu YD (2011) Ethanol outperforms multiple solvents in the extraction of chlorophyll-a from biological soil crusts. Soil Biol Biochem 43:857–861. https://doi.org/10.1016/j.soilbio.2010.12.007
Lan SB, Wu L, Zhang DL, Hu CX (2012) Composition of photosynthetic organisms and diurnal changes of photosynthetic efficiency in algae and moss crusts. Plant Soil 351:325–336. https://doi.org/10.1007/s11104-011-0966-9
Lan SB, Wu L, Zhang DL, Hu CX (2015) Effects of light and temperature on open cultivation of desert cyanobacterium Microcoleus vaginatus. Bioresource Technol 182:144–150. https://doi.org/10.1016/j.biortech.2015.02.002
Li XR, Chen YW, Yang LW (2004a) Cryptogam diversity and formation of soil crusts in temperate desert. Ann Arid Zone 43:335–353. https://www.researchgate.net/publication/287785714
Li XR, Xiao HL, Zhang JG, Wang XP (2004) Long-term ecosystem effects of sand-binding vegetation in the Tengger desert, Northern China. Restor Ecol 12:376–390. https://doi.org/10.1111/j.1061-2971.2004.00313.x
Li XR, Chen YW, Su YG, Tan HJ (2006) Effects of biological soil crust on desert insect diversity: evidence from the Tengger Desert of Northern China. Arid Land Res Manag 20:263–280. https://doi.org/10.1080/15324980600940985
Li XR, He MZ, Duan ZH, Xiao HL, Jia XH (2007) Recovery of topsoil physicochemical properties in revegetated sites in the sand-burial ecosystems of the Tengger Desert, northern China. Geomorphology 88:254–265. https://doi.org/10.1016/j.geomorph.2006.11.009
Li XR, He MZ, Zerbe S, Li XJ, Liu LC (2010) Micro-geomorphology determines community structure of biological soil crusts at small scales. Earth Surf Proc Land 35:932–940. https://doi.org/10.1002/esp.1963
Li XR, Jia RL, Chen YW, Huang L, Zhang P (2011) Association of ant nests with successional stages of biological soil crusts in the Tengger Desert, Northern China. Appl Soil Ecol 47:59–66. https://doi.org/10.1016/j.apsoil.2010.10.010
Li XR, Zhang ZS, Huang L, Wang XP (2013) Review of the ecohydrological processes and feedback mechanisms controlling sand-binding vegetation systems in sandy desert regions of China. Chinese Sci Bull 58:1483–1496. https://doi.org/10.1007/s11434-012-5662-5
Li JH, Li XR, Chen CY (2014) Degradation and reorganization of thylakoid protein complexes of Bryum argenteum in response to dehydration and rehydration. Bryologist 117:110–118. https://doi.org/10.1639/0007-2745-117.2.110
Li XR, Song G, Hui R, Zhang P (2017) Precipitation and topsoil attributes determine the species diversity and distribution patterns of crustal communities in desert ecosystems. Plant Soil 420:163–175. https://doi.org/10.1007/s11104-017-3385-8
Lichtenthaler HK, Wellburn AR (1983) Determinations of total carotenoids and chlorophylls a and b of leaf extracts in different solvents. Biochem Soc Trans 11:591–592. https://doi.org/10.1042/bst0110591
Liu JB, Feng W, Zhang YQ, Jia X, Wu B, Qin SG, Fa KY, Lai ZR (2015) Abiotic CO2 exchange between soil and atmosphere and its response to temperature. Environ Earth Sci 73:2463–2471. https://doi.org/10.1007/s12665-014-3595-9
Lu Y, Li XR, He MZ, Wang ZN, Tan HJ (2010) Nickel effects on growth and antioxidative enzymes activities in desert plant Zygophyllum xanthoxylon (Bunge) Maxim. Sci Cold Arid Reg 2:436–444
Maier S, Tamm A, Wu D, Caesar J, Grube M, Weber B (2018) Photoautotrophic organisms control microbial abundance, diversity, and physiology in different types of biological soil crusts. ISME J 12:1032–1046. https://doi.org/10.1038/s41396-018-0062-8
Oliver MJ, Bewley JD (2010) Desiccation-tolerance of plant tissues: a mechanistic overview. In: Janick J (ed) Horticultural Reviews, vol 18. Wiley, Hoboken, pp 171–213
Oliver MJ, Velten J, Mishler BD (2005) Desiccation tolerance in bryophytes: a reflection of the primitive strategy for plant survival in dehydrating habitats? Integr Comp Biol 45:788–799. https://doi.org/10.1093/icb/45.5.788
Pan Y, Wu LJ, Yu ZL (2006) Effect of salt and drought stress on antioxidant enzymes activities and SOD isoenzymes of liquorice (Glycyrrhiza uralensis Fisch). Plant Growth Regul 49:157–165. https://doi.org/10.1007/s10725-006-9101-y
Proctor MCF, Oliver MJ, Wood AJ, Alpert P, Stark LR, Cleavitt NL, Mishler BD (2007) Desiccation-tolerance in bryophytes: a review. Bryologist 110:595–621. https://doi.org/10.1639/0007-2745(2007)110[595:DIBAR]2.0.CO;2
Quan R, Shang M, Zhang H, Zhao Y, Zhang J (2004) Improved chilling tolerance by transformation with betA gene for the enhancement of glycine betaine synthesis in maize. Plant Sci 166:141–149. https://doi.org/10.1016/j.plantsci.2003.08.018
Ramiro DA, Guerreiro-Filho O, Mazzafera P (2006) Phenol contents, oxidase activities, and the resistance of coffee to the leaf miner Leucoptera coffeella. J Chem Ecol 32:1977–1988. https://doi.org/10.1007/s10886-006-9122-z
Reed SC, Coe KK, Sparks JP, Housman DC, Zelikova TJ, Belnap J (2012) Changes to dryland rainfall result in rapid moss mortality and altered soil fertility. Nat Clim Change 2:752–755. https://doi.org/10.1038/nclimate1596
Rivera-Aguilar V, Godinez-Alvarez H, Moreno-Torres R, Rodriguez-Zaragoza S (2009) Soil physico-chemical properties affecting the distribution of biological soil crusts along an environmental transect at Zapotitlan drylands, Mexico. J Arid Environ 73:1023–1028. https://doi.org/10.1016/j.jaridenv.2009.05.003
Roychoudhury A, Basu S (2012) Ascorbate-glutathione and plant tolerance to various abiotic stresses. In: Anjum NA, Umar S, Ahmad A (eds) Oxidative stress in plants: causes, consequences and tolerance. IK International, New Delhi, pp 177–258
Scandalios JG (1997) Oxidizing the genes oxidative stress and the molecular biology of antioxidant defences. Cold Spring Harbor Laboratory Press, New York, pp 383–388
Seel W, Hendry G, Atherton N, Lee J (1991) Radical formation and accumulation in vivo, in desiccation tolerant and intolerant mosses. Free Radic Res Commun 15:133–141. https://doi.org/10.3109/10715769109049133
Seel WE, Hendry GAF, Lee JA (1992) The combined effects of desiccation and irradiance on mosses from xeric and hydric habitats. J Exp Bot 43:1023–1030. https://doi.org/10.1093/jxb/43.8.1023
Shi LR, Liu ZH (2010) Influences of drought stress on antioxidative activity and osmoregulation substance of Sonchus brachyotus DC. Acta Agrestia Sinica 18:673–677. https://doi.org/10.3724/SP.J.1077.2010.01263
Stark LR (2005) Phenology of patch hydration, patch temperature and sexual reproductive output over a four-year period in the desert moss Crossidium crassinerve. J Bryol 27:231–240. https://doi.org/10.1179/174328205X69977
Tamm A, Caesar J, Kunz N, Colesie C, Reichenberger H, Weber B (2018) Ecophysiological properties of three biological soil crust types and their photoautotrophs from the Succulent Karoo, South Africa. Plant Soil 429:127–146. https://doi.org/10.1007/s11104-018-3635-4
Walter H, Stadelmann E (1968) The physiological prerequisites for the transition of autotrophic plants from water to terrestrial life. Bioscience 18:694–701. https://doi.org/10.2307/1294191
Wang XP, Kang ES, Zhang JG, Li XR (2004) Evapotranspiration of Artemisia ordosia vegetation in stabilized arid desert dune in Shapotou, China. Arid Land Res Manag 18:63–76. https://doi.org/10.1080/15324980490245022
Wang SK, Zuo XA, Zhao XY, Li YQ, Zhou X, Lv P, Luo YQ, Yun JY (2016) Responses of soil fungal community to the sandy grassland restoration in Horqin Sandy Land, northern China. Environ Monit Assess 188:21. https://doi.org/10.1007/s10661-015-5031-3
Xu GF, Zhang ZY (2009) Effect of high-temperature stress on physiological and biochemical indices of four Lysimachia plants. Chinese J Eco-Agri 17:565–569. https://doi.org/10.3724/SP.J.1011.2009.00565
Yu R, Zhang Z, Lu X, Chang IS, Liu T (2020) Variations in dew moisture regimes in desert ecosystems and their influencing factors. WIRS Water 7:e1482. https://doi.org/10.1002/wat2.1482
Zhang YM, Chen J, Wang L, Wang XQ, Gu ZH (2007) The spatial distribution patterns of biological soil crusts in the Gurbantunggut Desert, Northern Xinjiang, China. J Arid Environ 68:599–610. https://doi.org/10.1016/j.jaridenv.2006.06.012
Zhang J, Zhang YM, Downing A, Cheng JH, Zhou XB, Zhang BC (2009) The influence of biological soil crusts on dew deposition in Gurbantunggut Desert, Northwestern China. J Hydrol 379:220–228. https://doi.org/10.1016/j.jhydrol.2009.09.053
Zhang BC, Zhang YM, Downing A, Niu YL (2011) Distribution and composition of cyanobacteria and microalgae associated with biological soil crusts in the Gurbantunggut desert, China. Arid Land Res Manag 25:275–293. https://doi.org/10.1080/15324982.2011.565858
Zhang M, Hu CX, Zhao XH, Tan QL, Sun XC, Cao AY, Cui M, Zhang Y (2012) Molybdenum improves antioxidant and osmotic-adjustment ability against salt stress in Chinese cabbage (Brassica campestris L. ssp. Pekinensis). Plant Soil 355:375–383. https://doi.org/10.1007/s11104-011-1109-z
Zhang J, Zhang YM (2014) Diurnal variations of chlorophyll fluorescence and CO2 exchange of biological soil crusts in different successional stages in the Gurbantunggut Desert of northwestern China. Ecol Res 29:289–298. https://doi.org/10.1007/s11284-013-1122-1
Zhang ZS, Chen YL, Xu BX, Huang L, Tan HJ, Dong XJ (2015) Topographic differentiations of biological soil crusts and hydraulic properties in fixed sand dunes, Tengger Desert. J Arid Land 2:205–215. https://doi.org/10.1007/s40333-014-0048-y
Zhang BC, Li RH, Xiao P, Su YG, Zhang YM (2016) Cyanobacterial composition and spatial distribution based on pyrosequencing data in the Gurbantunggut Desert, Northwestern China. J Basic Microb 56:308–320. https://doi.org/10.1002/jobm.201500226
Zhang XQ, Zhao YZ, Wang SJ (2017) Responses of antioxidant defense system of epilithic mosses to drought stress in karst rock desertified areas. Acta Geochimica 36:205–212. https://doi.org/10.1007/s11631-017-0140-z
Zhang CP, Niu DC, Song ML, Elser JJ, Okie JG, Fu H (2018) Effects of rainfall manipulations on carbon exchange of cyanobacteria and moss-dominated biological soil crusts. Soil Biol Biochem 124:24–31. https://doi.org/10.1016/j.soilbio.2018.05.021
Zhao Y, Jia RL, Wang J (2019) Towards stopping land degradation in drylands: water-saving techniques for cultivating biocrusts in situ. Land Degrad Dev 30:2336–2346. https://doi.org/10.1002/ldr.3423
Zheng YP, Xu M, Zhao JC, Bei SQ, Hao LH (2011) Effects of inoculated Microcoleus vaginatus on the structure and function of biological soil crusts of desert. Biol Fert Soils 47:473–480. https://doi.org/10.1007/s00374-010-0521-5
Acknowledgements
The research was jointly supported by the National Key Research and Development Program of China (2018YFC0406603) and the National Natural Science Foundation of China (Nos. 41621001, 32061123006, and 41701104).
Author information
Authors and Affiliations
Contributions
All authors contributed to the study. Investigation, sample collection and measurement, data collection and analysis were performed by RH, HT, and RJ. XL and RZ carried out the manuscript revision. The first draft of the manuscript was written by RH and all authors commented on previous versions of the manuscript. All authors read and approved the final manuscript.
Corresponding author
Ethics declarations
Conflict of interest
The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.
Additional information
Responsible Editor: Manuel Delgado-Baquerizo.
Publisher's Note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Rights and permissions
About this article
Cite this article
Hui, R., Li, X., Zhao, R. et al. Physiological response of moss/cyanobacteria crusts along a precipitation gradient from semi-arid to arid desert in China. Plant Soil 468, 97–113 (2021). https://doi.org/10.1007/s11104-021-05117-2
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s11104-021-05117-2