青海祁连棱果沙棘自然杂交带三种沙棘光合特性日变化分析
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摘要
杂交带是研究物种形成及进化的天然实验室。为探究棱果沙棘自然杂交带三种沙棘的生理生态适应性,该研究在光照充足的7月中旬分别测定了该区域三种沙棘雌、雄株的光合特性及相关环境因子的日变化。结果表明:(1)同种沙棘雌、雄株间光合日变化规律基本一致,不同沙棘光合特性的日变化规律存在较大差异。中国沙棘(Hippophae rhamnoides ssp. sinensis)净光合速率(Pn)日变化呈明显的双峰曲线,峰值出现在光合有效辐射(PAR)和空气温度(Ta)相对较高的10:00和14:00,最高达到(19.53±5.35)μmol·m~(-2)·s~(-1);棱果沙棘(Hippophae goniocarpa)和肋果沙棘(H. neurocarpa ssp. neurocarpa) Pn日变化均呈近双峰曲线,在PAR和Ta相对较低的8:00和16:00有较高的值,最高值分别为(13.43±3.43)和(15.27±2.43)μmol·m~(-2)·s~(-1)。(2)三种沙棘水分利用效率(WUE)与Pn的日变化规律一致,但中国沙棘具有最高的WUE,达到(6.72±0.95)μmol·mmol~(-1),棱果沙棘和肋果沙棘的WUE日变化最高分别为(4.03±1.08)和(4.93±0.86)μmol·mmol~(-1)。(3)三种沙棘蒸腾速率(Tr)、气孔导度(Gs)、胞间二氧化碳浓度(Ci)和气孔限制值(Ls)的日变化规律相似,其中Gs均在10:00后一致下降,在12:00左右均出现不同程度的气孔关闭现象,这是其发生光合午休的主要原因之一。杂交带三种沙棘光合特性日变化差异主要表现在中国沙棘和其他两种沙棘之间,而杂交种棱果沙棘与亲本种肋果沙棘的变化规律基本一致,其Pn与WUE可能受到了Ta、PAR和空气相对湿度(RH)等环境因子的影响。
Hybrid zone is a natural laboratory for studying speciation and evolution. In order to explore the ecophysiological adaptability of three species of seabuchthorn( Hippophae),we measured the relevant environmental factors and diurnal changes in photosynthetic characteristics of female and male plants of the three Hippophae species in Qilian County of Qinghai Province in July. The results were as follows:( 1) The diurnal variation of photosynthesis between female and male plants of the same species was basically consistent,and there are differences in the diurnal variation of photosynthetic characteristics of the three Hippophae species. Diurnal changes of net photosynthetic rate( Pn) in H. rhamnoides ssp. sinensis showed a significant bimodal curve,with peak values at 10: 00 and 14: 00,and the highest value was( 19.53±5.35) μmol·m~(-2)·s~(-1) when photosynthetically active radiation( PAR) and air temperature( Ta). Although the diurnal changes of Pnin H. goniocarpa and H. neurocarpa ssp. neurocarpa were all nearly bimodal curve,the higher values at 8: 00 and 16: 00,and the highest values were( 13.43±3.43) and( 15.27±2.43) μmol·m~(-2)·s~(-1),repectively when PAR and Tawere lower.( 2) Water use efficiency( WUE) of the three species of Hippophae was consistent with the change of Pn,but H. rhamnoides ssp. sinensis had a higher WUE,which peaked at 10: 00 and 14: 00,and the highest value was( 6.7±0.95) μmol·mmol~(-1). The peak values of H. goniocarpa and H. neurocarpa ssp. neurocarpa were8: 00 and 16: 00,and the highest values were( 4.03±1.08) and( 4.93±0.86) μmol·mmol~(-1),respectively.( 3) The diurnal changes of transpiration rate( Tr),stomatal conductance( Gs),intercellular CO_2 concentration( Ci) and stomatal limiting value( Ls) of the three species of Hippophae were similar. Gsdecreased uniformly after 10: 00 among them,and there were different degrees of stomatal closure at around 12: 00,which was one of the causes of leaf photosynthetic depression in midday. In diurnal changes of photosynthesis,H. rhamnoides ssp. sinensis was significantly different from others,H. goniocarpa showed similar change trend with H. neurocarpa ssp. neurocarpa,and its Pnand WUE may be affected by environmental factors such as Ta,PAR and air relative humidity( RH).
Hybrid zone is a natural laboratory for studying speciation and evolution. In order to explore the ecophysiological adaptability of three species of seabuchthorn( Hippophae),we measured the relevant environmental factors and diurnal changes in photosynthetic characteristics of female and male plants of the three Hippophae species in Qilian County of Qinghai Province in July. The results were as follows:( 1) The diurnal variation of photosynthesis between female and male plants of the same species was basically consistent,and there are differences in the diurnal variation of photosynthetic characteristics of the three Hippophae species. Diurnal changes of net photosynthetic rate( Pn) in H. rhamnoides ssp. sinensis showed a significant bimodal curve,with peak values at 10: 00 and 14: 00,and the highest value was( 19.53±5.35) μmol·m~(-2)·s~(-1) when photosynthetically active radiation( PAR) and air temperature( Ta). Although the diurnal changes of Pnin H. goniocarpa and H. neurocarpa ssp. neurocarpa were all nearly bimodal curve,the higher values at 8: 00 and 16: 00,and the highest values were( 13.43±3.43) and( 15.27±2.43) μmol·m~(-2)·s~(-1),repectively when PAR and Tawere lower.( 2) Water use efficiency( WUE) of the three species of Hippophae was consistent with the change of Pn,but H. rhamnoides ssp. sinensis had a higher WUE,which peaked at 10: 00 and 14: 00,and the highest value was( 6.7±0.95) μmol·mmol~(-1). The peak values of H. goniocarpa and H. neurocarpa ssp. neurocarpa were8: 00 and 16: 00,and the highest values were( 4.03±1.08) and( 4.93±0.86) μmol·mmol~(-1),respectively.( 3) The diurnal changes of transpiration rate( Tr),stomatal conductance( Gs),intercellular CO_2 concentration( Ci) and stomatal limiting value( Ls) of the three species of Hippophae were similar. Gsdecreased uniformly after 10: 00 among them,and there were different degrees of stomatal closure at around 12: 00,which was one of the causes of leaf photosynthetic depression in midday. In diurnal changes of photosynthesis,H. rhamnoides ssp. sinensis was significantly different from others,H. goniocarpa showed similar change trend with H. neurocarpa ssp. neurocarpa,and its Pnand WUE may be affected by environmental factors such as Ta,PAR and air relative humidity( RH).
引文
ARNOLD ML,1996.Natural hybridization and introgression[M].Princeton:Princeton University Press.
ARNOLD ML,1997.Natural hybridization and evolution[M].Oxford,UK:Oxford University Press.
ARNTZ MA,DELPH LF,2001.Pattern and process:Evidence for the evolution of photosynthetic traits in natural populations[J].Oecologia,127(4):455-467.
BARTON NH,HEWITT GM,1985.Analysis of hybrid zones[J].Ann Rev Ecol Syst,16(1):113-148.
BARTON NH,2010.The role of hybridization in evolution[J].Mol Ecol,10(3):551-568.
CAMPBELL DR,GALEN C,WU CA,2005.Ecophysiology of first and second generation hybrids in a natural plant hybrid zone[J].Oecologia,144(2):214-225.
DONOVAN LA,EHLERINGER JR,1994.Potential for selection on plants for water-use efficiency as estimated by carbon isotope discrimination[J].Am J Bot,81(7):927-935.
DONOVAN LA,ROSENTHAL DR,SANCHEZ-VELENOSI M,et al.,2010.Are hybrid species more fit than ancestral parent species in the current hybrid species habitats?[J].JEvol Biol,23(4):805-816.
DU YJ,2008.The phenotype and molecular evidence of natural cross and hybrid zone between Hippophae rhamnoides ssp.sinensis and Hippophae neurocarpa ssp.neurocarpa[D].Lanzhou:Northwest Normal University.[杜玉娟,2008.中国沙棘与肋果沙棘间自然杂交和杂交带的表型及分子证据[D].兰州:西北师范大学.]
HEWITT GM,1988.Hybrid zones-Natural laboratories for evolutionary studies[J].Trends Ecol Evol,3(7):158-167.
JIANG LL,2013.The clone structure and clone reproductive fitness of hybrids and their parents of natural hybrid zone in Hippophae goniocarpa ssp.goniocarpa[D].Lanzhou:Northwest Normal University.[江龙龙,2013.棱果沙棘自然杂交带杂种和亲本居群的克隆结构和克隆繁育适合度比较研究[D].兰州:西北师范大学.]
JIANG W,JIANG WB,LI ZG,2007.Advance of researches on germplasm differences and genetic expression of photosynthetic traits in horticultural crops[J].Nonwood For Res,25(4):102-108.[姜武,姜卫兵,李志国,2007.园艺作物光合性状种质差异及遗传表现研究进展[J].经济林研究,25(4):102-108.]
JIANG YF,YAN R,SU X,et al.,2014.Molecular evidence for bidirectional hybrid origin and Hippophae rhamnoides ssp.sinensis as the mainly maternal plant of the diploid hybrid Hippophae goniocarpa(Elaeagnaceae)[J].Bull Bot Res,34(1):32-36.[蒋严妃,严容,苏雪,等,2014.二倍体杂交种棱果沙棘双向杂交起源及其母本主要来源于中国沙棘的分子证据[J].植物研究,34(1):32-36.]
JIN TT,FU BJ,LIU GH,et al.,2011.Diurnal changes of photosynthetic characteristics of Hippophae rhamnoides and the relevant environment factors at different slope locations[J].Acta Ecol Sin,31(7):1783-1793.[靳甜甜,傅伯杰,刘国华,等,2011.不同坡位沙棘光合日变化及其主要环境因子[J].生态学报,31(7):1783-1793.]
JOHNSTON JA,ARNOLD ML,DONOVAN LA,2003.High hybrid fitness at seed and seedling life history stages in Louisiana irises[J].J Ecol,91(3):438-446.
KIMBALL S,CAMPBELL D,2009.Physiological differences among two Penstemon species and their hybrids in field and common garden environments[J].New Phytol,181(2):478-488.
MOORE WS,1977.An evaluation of narrow hybrid zones in vertebrates[J].Quart Rev Biol,52(3):263-277.
SCHIERENBECK KA,MARSHALL JD,1993.Seasonal and diurnal patterns of photosynthetic gas-exchange for LoniceraSempervirens and L-Japonica(Caprifoliaceae)[J].Am JBot,80(11):1292-1299.
SCHWEITZER JA,MARTINSEN GD,WHITHAM TG,2002.Cottonwood hybrids gain fitness traits of both parents:A mechanism for their long-term persistence?[J].Am JBot,89(6):981-990.
SUN K,MA RJ,CHEN XL,et al.,2003.Hybrid origin of the diploid species Hippophae goniocarpa evidenced by the internal transcribed spacers(ITS)of nuclear rDNA[J].Belgian J Bot,136(1):91-96.
TANG JJ,2009.The fitness analysis of hybrids and parental species of Hippophae goniocarpa hybrid zone[D].Lanzhou:Northwest Normal University.[唐洁涓,2009.棱果沙棘自然杂交带亲本与杂交后代的适应性分析[D].兰州:西北师范大学.]
WANG AM,LIU ZX,ZU YG,2000.Daily course of photosynthesis in seedlings of natural birch[J].J Shanghai Univ,4(1):202-205.[王爱民,刘志学,祖元刚,2000.天然白桦种苗光合作用日进程[J].上海大学学报,4(1):202-205.]
WEI LY,ZENG DJ,ZHANG JL,et al.,2010.The photosynthetic characteristics of four forage grasses in karst rock desertification arease[J].Acta Pratac Sin,19(3):212-219.[韦兰英,曾丹娟,张建亮,等,2010.岩溶石漠化区四种牧草植物光合生理适应性特征[J].草业学报,19(3):212-219.]
YE ZP,HU WH,YAN XH,et al.,2016.Photosynthetic characteristics of different plant species based on a mechanistic model of light-response of photosynthesis[J].Chin J Ecol,35(9):2544-2552.[叶子飘,胡文海,闫小红,等,2016.基于光响应机理模型的不同植物光合特性[J].生态学杂志,35(9):2544-2552.]
ZHANG H,2005.Origin of isoploid hybridization of Hippophae goniocarpa:Evidence from chloroplast genomes and nuclear genomes[D].Lanzhou:Northwest Normal University.[张辉,2005.棱果沙棘的同倍体杂交起源:来自叶绿体基因组和核基因组的证据[D].兰州:西北师范大学.]
ZHAO S,HUANG QX,LI YL,et al.,2014.Effects of shading treatments on photosynthetic characteristics of Juniperus sabina Ant.seedlings[J].Acta Ecol Sin,34(8):1994-2002.[赵顺,黄秋娴,李玉灵,等,2014.遮荫处理对臭柏幼苗光合特性的影响[J].生态学报,34(8):1994-2002.]
ZHONG QC,WANG JT,ZHOU JH,et al.,2014.Effects of water table manipulation on leaf photosynthesis,morphology and growth of Phragmites australis and Imperata cylindrica in the reclaimed tidal wetland at Dongtan of Chongming Island,China[J].Chin J Appl Ecol,25(2):408-418.[仲启铖,王江涛,周剑虹,等,2014.水位调控对崇明东滩围垦区滩涂湿地芦苇和白茅光合、形态及生长的影响[J].应用生态学报,25(2):408-418.]
ZOU CM,WANG YQ,LIU Y,et al.,2015.Responses of photosynthesis and growth to weak light regime in four legume species[J].Chin J Plant Ecol,39(9):909-916.[邹长明,王允青,刘英,等,2015.四种豆科作物的光合生理和生长发育对弱光的响应[J].植物生态学报,39(9):909-916.]
ARNOLD ML,1997.Natural hybridization and evolution[M].Oxford,UK:Oxford University Press.
ARNTZ MA,DELPH LF,2001.Pattern and process:Evidence for the evolution of photosynthetic traits in natural populations[J].Oecologia,127(4):455-467.
BARTON NH,HEWITT GM,1985.Analysis of hybrid zones[J].Ann Rev Ecol Syst,16(1):113-148.
BARTON NH,2010.The role of hybridization in evolution[J].Mol Ecol,10(3):551-568.
CAMPBELL DR,GALEN C,WU CA,2005.Ecophysiology of first and second generation hybrids in a natural plant hybrid zone[J].Oecologia,144(2):214-225.
DONOVAN LA,EHLERINGER JR,1994.Potential for selection on plants for water-use efficiency as estimated by carbon isotope discrimination[J].Am J Bot,81(7):927-935.
DONOVAN LA,ROSENTHAL DR,SANCHEZ-VELENOSI M,et al.,2010.Are hybrid species more fit than ancestral parent species in the current hybrid species habitats?[J].JEvol Biol,23(4):805-816.
DU YJ,2008.The phenotype and molecular evidence of natural cross and hybrid zone between Hippophae rhamnoides ssp.sinensis and Hippophae neurocarpa ssp.neurocarpa[D].Lanzhou:Northwest Normal University.[杜玉娟,2008.中国沙棘与肋果沙棘间自然杂交和杂交带的表型及分子证据[D].兰州:西北师范大学.]
HEWITT GM,1988.Hybrid zones-Natural laboratories for evolutionary studies[J].Trends Ecol Evol,3(7):158-167.
JIANG LL,2013.The clone structure and clone reproductive fitness of hybrids and their parents of natural hybrid zone in Hippophae goniocarpa ssp.goniocarpa[D].Lanzhou:Northwest Normal University.[江龙龙,2013.棱果沙棘自然杂交带杂种和亲本居群的克隆结构和克隆繁育适合度比较研究[D].兰州:西北师范大学.]
JIANG W,JIANG WB,LI ZG,2007.Advance of researches on germplasm differences and genetic expression of photosynthetic traits in horticultural crops[J].Nonwood For Res,25(4):102-108.[姜武,姜卫兵,李志国,2007.园艺作物光合性状种质差异及遗传表现研究进展[J].经济林研究,25(4):102-108.]
JIANG YF,YAN R,SU X,et al.,2014.Molecular evidence for bidirectional hybrid origin and Hippophae rhamnoides ssp.sinensis as the mainly maternal plant of the diploid hybrid Hippophae goniocarpa(Elaeagnaceae)[J].Bull Bot Res,34(1):32-36.[蒋严妃,严容,苏雪,等,2014.二倍体杂交种棱果沙棘双向杂交起源及其母本主要来源于中国沙棘的分子证据[J].植物研究,34(1):32-36.]
JIN TT,FU BJ,LIU GH,et al.,2011.Diurnal changes of photosynthetic characteristics of Hippophae rhamnoides and the relevant environment factors at different slope locations[J].Acta Ecol Sin,31(7):1783-1793.[靳甜甜,傅伯杰,刘国华,等,2011.不同坡位沙棘光合日变化及其主要环境因子[J].生态学报,31(7):1783-1793.]
JOHNSTON JA,ARNOLD ML,DONOVAN LA,2003.High hybrid fitness at seed and seedling life history stages in Louisiana irises[J].J Ecol,91(3):438-446.
KIMBALL S,CAMPBELL D,2009.Physiological differences among two Penstemon species and their hybrids in field and common garden environments[J].New Phytol,181(2):478-488.
MOORE WS,1977.An evaluation of narrow hybrid zones in vertebrates[J].Quart Rev Biol,52(3):263-277.
SCHIERENBECK KA,MARSHALL JD,1993.Seasonal and diurnal patterns of photosynthetic gas-exchange for LoniceraSempervirens and L-Japonica(Caprifoliaceae)[J].Am JBot,80(11):1292-1299.
SCHWEITZER JA,MARTINSEN GD,WHITHAM TG,2002.Cottonwood hybrids gain fitness traits of both parents:A mechanism for their long-term persistence?[J].Am JBot,89(6):981-990.
SUN K,MA RJ,CHEN XL,et al.,2003.Hybrid origin of the diploid species Hippophae goniocarpa evidenced by the internal transcribed spacers(ITS)of nuclear rDNA[J].Belgian J Bot,136(1):91-96.
TANG JJ,2009.The fitness analysis of hybrids and parental species of Hippophae goniocarpa hybrid zone[D].Lanzhou:Northwest Normal University.[唐洁涓,2009.棱果沙棘自然杂交带亲本与杂交后代的适应性分析[D].兰州:西北师范大学.]
WANG AM,LIU ZX,ZU YG,2000.Daily course of photosynthesis in seedlings of natural birch[J].J Shanghai Univ,4(1):202-205.[王爱民,刘志学,祖元刚,2000.天然白桦种苗光合作用日进程[J].上海大学学报,4(1):202-205.]
WEI LY,ZENG DJ,ZHANG JL,et al.,2010.The photosynthetic characteristics of four forage grasses in karst rock desertification arease[J].Acta Pratac Sin,19(3):212-219.[韦兰英,曾丹娟,张建亮,等,2010.岩溶石漠化区四种牧草植物光合生理适应性特征[J].草业学报,19(3):212-219.]
YE ZP,HU WH,YAN XH,et al.,2016.Photosynthetic characteristics of different plant species based on a mechanistic model of light-response of photosynthesis[J].Chin J Ecol,35(9):2544-2552.[叶子飘,胡文海,闫小红,等,2016.基于光响应机理模型的不同植物光合特性[J].生态学杂志,35(9):2544-2552.]
ZHANG H,2005.Origin of isoploid hybridization of Hippophae goniocarpa:Evidence from chloroplast genomes and nuclear genomes[D].Lanzhou:Northwest Normal University.[张辉,2005.棱果沙棘的同倍体杂交起源:来自叶绿体基因组和核基因组的证据[D].兰州:西北师范大学.]
ZHAO S,HUANG QX,LI YL,et al.,2014.Effects of shading treatments on photosynthetic characteristics of Juniperus sabina Ant.seedlings[J].Acta Ecol Sin,34(8):1994-2002.[赵顺,黄秋娴,李玉灵,等,2014.遮荫处理对臭柏幼苗光合特性的影响[J].生态学报,34(8):1994-2002.]
ZHONG QC,WANG JT,ZHOU JH,et al.,2014.Effects of water table manipulation on leaf photosynthesis,morphology and growth of Phragmites australis and Imperata cylindrica in the reclaimed tidal wetland at Dongtan of Chongming Island,China[J].Chin J Appl Ecol,25(2):408-418.[仲启铖,王江涛,周剑虹,等,2014.水位调控对崇明东滩围垦区滩涂湿地芦苇和白茅光合、形态及生长的影响[J].应用生态学报,25(2):408-418.]
ZOU CM,WANG YQ,LIU Y,et al.,2015.Responses of photosynthesis and growth to weak light regime in four legume species[J].Chin J Plant Ecol,39(9):909-916.[邹长明,王允青,刘英,等,2015.四种豆科作物的光合生理和生长发育对弱光的响应[J].植物生态学报,39(9):909-916.]