喀什霸王的结实和种子萌发特性
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摘要
喀什霸王(Zygophyllum kaschgaricum)是生长于中国新疆南部荒漠环境的稀有种及二级保护植物。当前,该物种在自然种群中呈分散式及片段化分布,且种群密度低,种群老龄化较严重。因此,为了了解该物种种子萌发特性及其对荒漠环境的响应,该文采用室内控制实验方法,对该物种的自然坐果率、结籽率、种子吸水特性、种子休眠和萌发特性及对干旱胁迫的响应进行了比较研究。结果表明:喀什霸王在自然种群中的坐果率及结籽率较低,且种子的败育率较高。不同干藏时间种子的吸水速率间存在显著差异;随着干藏时间的延长,种子的吸水率逐步增强。刚成熟的种子在不同温度及光周期下均可萌发;其中高温(10/20℃, 20/30℃)及黑暗条件下的萌发率比低温(10/5℃, 5/2℃)及光照条件下的萌发率高。不同干藏时间的种子在不同浓度赤霉素(GA3)下的萌发率均较高;但低温储藏时间对该物种种子的打破休眠及萌发率没有促进作用。以上结果说明该物种存在非深度生理休眠;而干藏时间、高温且黑暗及高浓度(50 mmol·L–1) GA3是打破休眠及促进种子萌发的最合适条件。高温条件下的干旱胁迫对喀什霸王种子萌发具有抑制作用;春季和秋季降水量决定种子的萌发率。总之,喀什霸王种子在物候上表现出的春秋季萌动及非深度生理休眠以提高幼苗存活力及保障种群稳定性,是一种对新疆南部干旱及高温胁迫荒漠环境的适应策略。
Aims Zygophyllum kaschgaricum is a rare and secondary protection plant, which grows in the desert environment in southern Xinjiang, China. At present, this species not only distributes decentralized and fragmentally, but also is low population density and severe ageing of populations in the natural populations. To better understand how this species is adapted to its desert habitat, we studied seed set and its germination characteristics and addressed the following questions. 1) What are the seed set characteristics and water absorption of seeds in different dry storage duration in natural population? 2) What are the seed dormancy and drought stress responses of this species in desert environment? Methods For the seed set and seed germination characters, we compared the natural seed set of each fruit and water absorption characteristics of each seed in the different dry storage duration, seed dormancy and germination and their responses to drought stress, and the adaptive strategies of this species in southern Xinjiang(Taklimakan desert) natural populations were analyzed. Important findings The natural fruit and seed set rate of this species is low and seed abortion rate is high in natural populations. There is significant difference in the water absorption capacity of seeds subjected to different durations of dry storage time(p < 0.001), and water absorption capacity of seeds increases with the extension of dry storage time. The seed germination rate increased with the extension of dry storage time under different temperature conditions. The newly matured seeds germinate under different temperature conditions and light cycle(dark 12 h and light 12 h); The germination rate at high temperature(10/20 ℃, 20/30 ℃) and dark conditions is higher than that at low temperature(10/5 ℃, 5/2 ℃) and light conditions. Seed germination rate at different light and temperature conditions under different GA3 concentration was high, but storage period at low temperature does not promote breaking of dormancy and germination in Z. kaschgaricum seeds. The results suggest that the species has shallow physiological dormancy. Storage at dry environment, high temperature and dark mode conditions, and high concentration(50 mmol·L–1) of GA3 were the most suitable factors to break dormancy and thus promote seed germination of Z. kaschgaricum. Drought stress under the high temperature inhibited seed germination. The rainfall during spring and autumn is the most important factor to control the seed germination rate. However, shallow physiological dormancy and seed germination at spring and autumn characteristics can improve for the seedling viability and population expansion of this Z. kaschgaricum in the desert, which may be an adaptive strategy to drought and high temperature stress of desert environment in Taklimakan(southern Xinjiang), China.
Aims Zygophyllum kaschgaricum is a rare and secondary protection plant, which grows in the desert environment in southern Xinjiang, China. At present, this species not only distributes decentralized and fragmentally, but also is low population density and severe ageing of populations in the natural populations. To better understand how this species is adapted to its desert habitat, we studied seed set and its germination characteristics and addressed the following questions. 1) What are the seed set characteristics and water absorption of seeds in different dry storage duration in natural population? 2) What are the seed dormancy and drought stress responses of this species in desert environment? Methods For the seed set and seed germination characters, we compared the natural seed set of each fruit and water absorption characteristics of each seed in the different dry storage duration, seed dormancy and germination and their responses to drought stress, and the adaptive strategies of this species in southern Xinjiang(Taklimakan desert) natural populations were analyzed. Important findings The natural fruit and seed set rate of this species is low and seed abortion rate is high in natural populations. There is significant difference in the water absorption capacity of seeds subjected to different durations of dry storage time(p < 0.001), and water absorption capacity of seeds increases with the extension of dry storage time. The seed germination rate increased with the extension of dry storage time under different temperature conditions. The newly matured seeds germinate under different temperature conditions and light cycle(dark 12 h and light 12 h); The germination rate at high temperature(10/20 ℃, 20/30 ℃) and dark conditions is higher than that at low temperature(10/5 ℃, 5/2 ℃) and light conditions. Seed germination rate at different light and temperature conditions under different GA3 concentration was high, but storage period at low temperature does not promote breaking of dormancy and germination in Z. kaschgaricum seeds. The results suggest that the species has shallow physiological dormancy. Storage at dry environment, high temperature and dark mode conditions, and high concentration(50 mmol·L–1) of GA3 were the most suitable factors to break dormancy and thus promote seed germination of Z. kaschgaricum. Drought stress under the high temperature inhibited seed germination. The rainfall during spring and autumn is the most important factor to control the seed germination rate. However, shallow physiological dormancy and seed germination at spring and autumn characteristics can improve for the seedling viability and population expansion of this Z. kaschgaricum in the desert, which may be an adaptive strategy to drought and high temperature stress of desert environment in Taklimakan(southern Xinjiang), China.
引文
Ahmad S,Ahmad R,Ashraf MY,Ashraf M,Waraich EA(2009).Sunflower(Helianthus annuus L.)response to drought stress at germination and seedling growth stages.Pakistan Journal of Botany,41,647-654.
Albrecht MA,McCarthy BC(2011).Variation in dormancy and germination in three co-occurring perennial forest herbs.Plant Ecology,212,1465-1477.
Baskin CC,Baskin JM(1988).Germination ecophysiology of herbaceous plant species in a temperate region.American Journal of Botany,75,286-305.
Baskin CC,Baskin JM(2014).Seed:Ecology,Biogeography,and Evolution of Dormancy and Germination.Academic Press,San Diego,USA.215-375.
Baskin JM,Baskin CC(2004).A classification system for seed dormancy.Seed Science Research,14,1-16.
Brown AR,Owena SF,Petersa J(2015).Climate change and pollution speed declines in zebra fish populations.Proceedings of the National Academy of the Sciences of the United States of America,112,1237-1246.
Casas RR,Kovach K,Dittmar E(2012).Seed after-ripening and dormancy determine adult life history independently of germination timing.New Phytologist,194,868-879.
Copete MA,Herranz JM,Ferrandis P(2015).Annual dormancy cycles in buried seeds of shrub species:Germination ecology of Sideritis serrata(Labiatae).Plant Biology,17,798-807.
Galloway LF,Etterson JR(2009).Plasticity to canopy shade in a monocarpic herb:Within-and between-generation effects.New Phytologist,182,1003-1012.
Gutterman Y(1993).Seed Germination in Desert Plants.Springer-Verlag,Berlin Heidelberg.
Gutterman Y(2000).Environmental factors and survival strategies of annual plant species in the Negev Desert,Israel.Plant Species Biology,15,113-125.
Huang ZY,Gutterman Y,Hu ZH,Zhang XS(2001).Seed germination in Artemisia sphaerocephala II.The influencing of environmental factors.Acta Phytoecologica Sinica,25,240-246.[黄振英,Gutterman Y,胡正海,张新时(2001).白沙蒿种子萌发特性的研究II.环境因素的影响.植物生态学报,25,240-246.]
Kalisz S,Wardle GM(1994).Life history variation in Campanula americana(Campanulaceae):Population differentiation.American Journal of Botany,81,521-527.
Leadley P,Pereira HM,Alkemade R(2010).Biodiversity Scenarios:Projections of 21st Century Change in Biodiversity and Associated Ecosystem Services.Secretariat of the Convention on Biological Diversity,Montreal.121-133.
Liu B,LüXG,Jiang M,Zhang WG,Wu HT(2015).Effects of light and water depth on seed germination of Phragmites australis in the wetlands of Songnen Plain.Chinese Journal of Plant Ecology,39,616-620.[刘波,吕宪国,姜明,张文广,武海涛(2015).光照、水深交互作用对松嫩湿地芦苇种子萌发的影响.植物生态学报,39,616-620.]
Lu JJ,Tan DY,Baskin JM,Baskin CC(2014).Germination season and watering regime,but not seed morph,affect life history traits in a cold desert diaspore-heteromorphic annual.PLOS ONE,9,e102018.DOI:10.1371/journal.pone.0102018.
Lu JJ,Zhou YM,Tan DY,Baskin CC,Baskin JM(2015).Seed dormancy in six cold desert Brassicaceae species with indehiscent fruits.Seed Science Research,25,276-285.
Mandák B(2001).Germination requirements of invasive and non-invasive Atriplex species:A comparative study.Flora,198,45-54.
Neaves LE,Eales J,Whitlock R(2015).The fitness consequences of inbreeding in natural populations and their implications for species conservation-A systematic map.Environmental Evidence,4,2-17.
Nordborg M,Bergelson J(1999).The effect of seed and rosette cold treatment on germination and flowering time in some Arabidopsis thaliana(Brassicaceae)ecotypes.American Journal of Botany,86,470-475.
Nurulla M,Baskin CC,Lu JJ,Tan DY,Baskin JM(2015).Intermediate morphophysiological dormancy allows for life-cycle diversity in the annual weed,Turgenia latifolia(Apiaceae).Australian Journal of Botany,62,630-637.
Parmesan C,Yohe G(2003).A globally coherent fingerprint of climate change impacts across natural systems.Nature,421,37-42.
Pluntz M,Coz SL,Peyrard N,Pradel R,Choquet R,Cheptou PO(2018).A general method for estimating seed dormancy and colonisation in annual plants from the observation of existing flora.Ecology Letters,21,1311-1318.
Qin HN,Zhao LN,Yu SX,Liu HY,Liu B,Xia NH,Peng H,Li ZY,Zhang ZX,He XJ,Yin LK,Lin YL,Liu QR,Hou YT,Liu Y,Liu QX,Cao W,Li JQ,Chen SL,Jin XH,Gao TG,Chen WL,Ma HY,Geng YY,Jin XF,Chang CY,Jiang H,Cai L,Zang CX,Wu JY,Ye JF,Lai YJ,Liu B,Lin QW,Xue NX(2017).Evaluating the endangerment status of China’s angiosperms through the red list assessment.Biodiversity Science,25,745-757.[覃海宁,赵莉娜,于胜祥,刘慧圆,刘博,夏念和,彭华,李振宇,张志翔,何兴金,尹林克,林余霖,刘全儒,侯元同,刘演,刘启新,曹伟,李建强,陈世龙,金效华,高天刚,陈文俐,马海英,耿玉英,金孝锋,常朝阳,蒋宏,蔡蕾,臧春鑫,武建勇,叶建飞,赖阳均,刘冰,林秦文,薛纳新(2017).中国被子植物濒危等级的评估.生物多样性,25,745-757.]
Ren H,Zhang QM,Lu HF,Liu HX,Guo QF,Wang J,Jian SG,Bao HO(2012).Wild plant species with extremely small populations require conservation and reintroduction in China.AMBIO,41,913-917.
Soltani E,Gruber S,Oveisi M,Salehi N,Alahdadi I,Javid MG(2017).Water stress,temperature regimes and light control induction,and loss of secondary dormancy in Brassica napus L.seeds.Seed Science Research,27,217-230.
Song NP,Wang X,Chen L,Xue Y,Chen J,Sui JM,Wang L,Yang XG(2018).Co-existence mechanisms of plant species within“soil islands”habitat of desert steppe.Biodiversity Science,26,667-677.[宋乃平,王兴,陈林,薛毅,陈娟,随金明,王磊,杨新国(2018).荒漠草原“土岛”生境群落物种共存机制.生物多样性,26,667-677.]
Sun HZ,Lu JJ,Tan DY,Baskin JM,Baskin CC(2009).Dormancy and germination characteristics of the trimorphic achenes of Garhadiolus papposus(Asteraceae),an annual ephemeral from the Junggar Desert,China.South African Journal of Botany,75,537-545.
Sun QL,Zhang RH,Yi SG,Yang L,Li QY,Zhou JH,Lai LM,Jiang LH,Zheng YR(2019).Responses of germination and seedling emergence of Stipa bungeana to key environmental factors.Acta Ecologica Sinica,39,2034-2042.[孙清琳,张瑞红,易三桂,杨柳,李巧燕,周继华,来利明,姜联合,郑元润(2019).长芒草(Stipa bungeana)种子萌发与出苗对关键环境因子的响应.生态学报,39,2034-2042.]
Tobe K,Zhang L,Yu QG(2001).Characteristics of seed germination in five non-halophytic Chinese desert shrub species.Journal of Arid Environments,47,191-201.
Venable DL,Búrquez A,Corral G,Morales E,Espinosa F(1987).The ecology of seed heteromorphism in Heterosperma pinnatum in central Mexico.Ecology,68,65-76.
Western D(2001).Human-modified ecosystems and future evolution.Proceedings of the National Academy of Sciences of United States of America,98,5458-5465.
Xia Q,Maharajah P,Cueff G,Rajjouc L,Prodhommed D,Gibond Y,Baillya C,Corbineaua F,Meimouna P,El-Maarouf-Bouteau H(2018).Integrating proteomics and enzymatic profiling to decipher seed metabolism affected by temperature in seed dormancy and germination.Plant Science,269,118-125.
Yao H,Tan DY(2005).Size-dependent reproductive output and life-history strategies in four ephemeral species of Trigonella.Acta Phytoecologica Sinica,29,954-960.[姚红,谭敦炎(2005).胡卢巴属4种短命植物个体大小依赖的繁殖输出与生活史对策.植物生态学报,29,954-960.]
Yin LK(2006).Rare,Endangered and Endemic Higher Plants in Xinjiang.Xinjiang Science and Technology Press,ürümqi.35-45.[尹林克(2006).新疆珍稀濒危特有高等植物.新疆科学技术出版社,乌鲁木齐.35-45.]
Zhang WH,Xu XB,Zhou JY(2004).Distribution and bioecological characteristics of Abies chensiensis,an endangered plant.Chinese Biodiversity,12,419-426.[张文辉,许晓波,周建云(2004).濒危植物秦岭冷杉地理分布和生物生态学特性研究.生物多样性,12,419-426.]
Albrecht MA,McCarthy BC(2011).Variation in dormancy and germination in three co-occurring perennial forest herbs.Plant Ecology,212,1465-1477.
Baskin CC,Baskin JM(1988).Germination ecophysiology of herbaceous plant species in a temperate region.American Journal of Botany,75,286-305.
Baskin CC,Baskin JM(2014).Seed:Ecology,Biogeography,and Evolution of Dormancy and Germination.Academic Press,San Diego,USA.215-375.
Baskin JM,Baskin CC(2004).A classification system for seed dormancy.Seed Science Research,14,1-16.
Brown AR,Owena SF,Petersa J(2015).Climate change and pollution speed declines in zebra fish populations.Proceedings of the National Academy of the Sciences of the United States of America,112,1237-1246.
Casas RR,Kovach K,Dittmar E(2012).Seed after-ripening and dormancy determine adult life history independently of germination timing.New Phytologist,194,868-879.
Copete MA,Herranz JM,Ferrandis P(2015).Annual dormancy cycles in buried seeds of shrub species:Germination ecology of Sideritis serrata(Labiatae).Plant Biology,17,798-807.
Galloway LF,Etterson JR(2009).Plasticity to canopy shade in a monocarpic herb:Within-and between-generation effects.New Phytologist,182,1003-1012.
Gutterman Y(1993).Seed Germination in Desert Plants.Springer-Verlag,Berlin Heidelberg.
Gutterman Y(2000).Environmental factors and survival strategies of annual plant species in the Negev Desert,Israel.Plant Species Biology,15,113-125.
Huang ZY,Gutterman Y,Hu ZH,Zhang XS(2001).Seed germination in Artemisia sphaerocephala II.The influencing of environmental factors.Acta Phytoecologica Sinica,25,240-246.[黄振英,Gutterman Y,胡正海,张新时(2001).白沙蒿种子萌发特性的研究II.环境因素的影响.植物生态学报,25,240-246.]
Kalisz S,Wardle GM(1994).Life history variation in Campanula americana(Campanulaceae):Population differentiation.American Journal of Botany,81,521-527.
Leadley P,Pereira HM,Alkemade R(2010).Biodiversity Scenarios:Projections of 21st Century Change in Biodiversity and Associated Ecosystem Services.Secretariat of the Convention on Biological Diversity,Montreal.121-133.
Liu B,LüXG,Jiang M,Zhang WG,Wu HT(2015).Effects of light and water depth on seed germination of Phragmites australis in the wetlands of Songnen Plain.Chinese Journal of Plant Ecology,39,616-620.[刘波,吕宪国,姜明,张文广,武海涛(2015).光照、水深交互作用对松嫩湿地芦苇种子萌发的影响.植物生态学报,39,616-620.]
Lu JJ,Tan DY,Baskin JM,Baskin CC(2014).Germination season and watering regime,but not seed morph,affect life history traits in a cold desert diaspore-heteromorphic annual.PLOS ONE,9,e102018.DOI:10.1371/journal.pone.0102018.
Lu JJ,Zhou YM,Tan DY,Baskin CC,Baskin JM(2015).Seed dormancy in six cold desert Brassicaceae species with indehiscent fruits.Seed Science Research,25,276-285.
Mandák B(2001).Germination requirements of invasive and non-invasive Atriplex species:A comparative study.Flora,198,45-54.
Neaves LE,Eales J,Whitlock R(2015).The fitness consequences of inbreeding in natural populations and their implications for species conservation-A systematic map.Environmental Evidence,4,2-17.
Nordborg M,Bergelson J(1999).The effect of seed and rosette cold treatment on germination and flowering time in some Arabidopsis thaliana(Brassicaceae)ecotypes.American Journal of Botany,86,470-475.
Nurulla M,Baskin CC,Lu JJ,Tan DY,Baskin JM(2015).Intermediate morphophysiological dormancy allows for life-cycle diversity in the annual weed,Turgenia latifolia(Apiaceae).Australian Journal of Botany,62,630-637.
Parmesan C,Yohe G(2003).A globally coherent fingerprint of climate change impacts across natural systems.Nature,421,37-42.
Pluntz M,Coz SL,Peyrard N,Pradel R,Choquet R,Cheptou PO(2018).A general method for estimating seed dormancy and colonisation in annual plants from the observation of existing flora.Ecology Letters,21,1311-1318.
Qin HN,Zhao LN,Yu SX,Liu HY,Liu B,Xia NH,Peng H,Li ZY,Zhang ZX,He XJ,Yin LK,Lin YL,Liu QR,Hou YT,Liu Y,Liu QX,Cao W,Li JQ,Chen SL,Jin XH,Gao TG,Chen WL,Ma HY,Geng YY,Jin XF,Chang CY,Jiang H,Cai L,Zang CX,Wu JY,Ye JF,Lai YJ,Liu B,Lin QW,Xue NX(2017).Evaluating the endangerment status of China’s angiosperms through the red list assessment.Biodiversity Science,25,745-757.[覃海宁,赵莉娜,于胜祥,刘慧圆,刘博,夏念和,彭华,李振宇,张志翔,何兴金,尹林克,林余霖,刘全儒,侯元同,刘演,刘启新,曹伟,李建强,陈世龙,金效华,高天刚,陈文俐,马海英,耿玉英,金孝锋,常朝阳,蒋宏,蔡蕾,臧春鑫,武建勇,叶建飞,赖阳均,刘冰,林秦文,薛纳新(2017).中国被子植物濒危等级的评估.生物多样性,25,745-757.]
Ren H,Zhang QM,Lu HF,Liu HX,Guo QF,Wang J,Jian SG,Bao HO(2012).Wild plant species with extremely small populations require conservation and reintroduction in China.AMBIO,41,913-917.
Soltani E,Gruber S,Oveisi M,Salehi N,Alahdadi I,Javid MG(2017).Water stress,temperature regimes and light control induction,and loss of secondary dormancy in Brassica napus L.seeds.Seed Science Research,27,217-230.
Song NP,Wang X,Chen L,Xue Y,Chen J,Sui JM,Wang L,Yang XG(2018).Co-existence mechanisms of plant species within“soil islands”habitat of desert steppe.Biodiversity Science,26,667-677.[宋乃平,王兴,陈林,薛毅,陈娟,随金明,王磊,杨新国(2018).荒漠草原“土岛”生境群落物种共存机制.生物多样性,26,667-677.]
Sun HZ,Lu JJ,Tan DY,Baskin JM,Baskin CC(2009).Dormancy and germination characteristics of the trimorphic achenes of Garhadiolus papposus(Asteraceae),an annual ephemeral from the Junggar Desert,China.South African Journal of Botany,75,537-545.
Sun QL,Zhang RH,Yi SG,Yang L,Li QY,Zhou JH,Lai LM,Jiang LH,Zheng YR(2019).Responses of germination and seedling emergence of Stipa bungeana to key environmental factors.Acta Ecologica Sinica,39,2034-2042.[孙清琳,张瑞红,易三桂,杨柳,李巧燕,周继华,来利明,姜联合,郑元润(2019).长芒草(Stipa bungeana)种子萌发与出苗对关键环境因子的响应.生态学报,39,2034-2042.]
Tobe K,Zhang L,Yu QG(2001).Characteristics of seed germination in five non-halophytic Chinese desert shrub species.Journal of Arid Environments,47,191-201.
Venable DL,Búrquez A,Corral G,Morales E,Espinosa F(1987).The ecology of seed heteromorphism in Heterosperma pinnatum in central Mexico.Ecology,68,65-76.
Western D(2001).Human-modified ecosystems and future evolution.Proceedings of the National Academy of Sciences of United States of America,98,5458-5465.
Xia Q,Maharajah P,Cueff G,Rajjouc L,Prodhommed D,Gibond Y,Baillya C,Corbineaua F,Meimouna P,El-Maarouf-Bouteau H(2018).Integrating proteomics and enzymatic profiling to decipher seed metabolism affected by temperature in seed dormancy and germination.Plant Science,269,118-125.
Yao H,Tan DY(2005).Size-dependent reproductive output and life-history strategies in four ephemeral species of Trigonella.Acta Phytoecologica Sinica,29,954-960.[姚红,谭敦炎(2005).胡卢巴属4种短命植物个体大小依赖的繁殖输出与生活史对策.植物生态学报,29,954-960.]
Yin LK(2006).Rare,Endangered and Endemic Higher Plants in Xinjiang.Xinjiang Science and Technology Press,ürümqi.35-45.[尹林克(2006).新疆珍稀濒危特有高等植物.新疆科学技术出版社,乌鲁木齐.35-45.]
Zhang WH,Xu XB,Zhou JY(2004).Distribution and bioecological characteristics of Abies chensiensis,an endangered plant.Chinese Biodiversity,12,419-426.[张文辉,许晓波,周建云(2004).濒危植物秦岭冷杉地理分布和生物生态学特性研究.生物多样性,12,419-426.]
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