伊犁郁金香的繁殖生物学特性及其生态适应对策
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摘要
准噶尔荒漠早春环境严酷多变，开花的植物种类少，而传粉昆虫的种类较少且其数量和质量在时空上的变化较大。生长在早春环境的植物，通过其花部结构、交配系统及传粉者间关系以及繁殖对策来适应恶劣的不良环境。因此，研究早春开花植物的繁殖生物学特性对于揭示植物与传粉者的关系及其对早春环境的适应具有重要科学意义。伊犁郁金香(Tulipa iliensis Regel.)是中亚地区及我国新疆北部分布的早春植物。该物种的花存在暂时性闭合现象，在自然居群的个体大小以及居群不同时期的开放花密度和传粉者数量间存在差异。本论文通过野外和室内的控制性试验，对伊犁郁金香的花部综合征及传粉特性与交配系统的关系、花暂时性闭合对繁殖成功的贡献、传粉者和花粉传递效率的时间性变化对结籽率的影响、花期依赖的花粉限制和近交衰退指数以及花大小依赖的性分配特点与传粉特性和交配系统的关系进行了研究，以期揭示该物种的繁殖特性及其对早春环境的生态适应对策。主要结果如下：
1.花部综合征和传粉特性对交配方式的影响及其在早春环境的适应进行了研究。该物种花具有温度变化导致的花暂时性闭合特性、花不完全雌雄异位、花内雌雄同熟及花开放持续时间的花龄依赖的变化等花部综合征；花瓣、雄蕊及雌蕊均能分泌香味。花蜜体积及传粉者访花频率随着花龄的增大而逐渐减少；其访花频率与花粉活力、柱头可授性及花蜜体积间均存在正相关关系。自花授粉与异花授粉花均结实，属于混合交配系统；存在自主自交及意蜂(Apis mellifera)在花上的运动模式引起的协助自交。不同处理的花所产生的种子萌发率间均存在显著差异；异花授粉种子的适合度比自然授粉和自花授粉种子高。花雌雄功能分离的时空特点、花报酬及花暂时性闭合等花部综合征有关的自主自交及传粉者行为导致的协助自交是该物种在长期适应传粉者限制的早春环境中，为保障繁殖成功而形成的环境适应对策。
2.花暂时性闭合对繁殖成功的贡献进行了野外和室内的控制性试验。在花完全开放和完全闭合期间的柱头和花药空间位置间存在差异；花暂时性闭合能促进自主自交。花暂时性闭合能保持花内微环境温度，提供较稳定的花微环境温度来影响花粉萌发率及柱头可授性。在下雨前伊犁郁金香通过花暂时性闭合来避免雨水对花药上花粉的冲刷以及对花粉活力和柱头可授性的影响。在大风前的花暂时性通过保护花粉聚集水平来影响传粉者的传粉效率或促进自主自交率。伊犁郁金香花的暂时性闭合是该物种在长期适应早春环境中，为了促进自主自交、延长雌雄器官的功能及提高繁殖成功而进化出的非花部结构机制。
3.在自然居群中，传粉者及花粉传递效率的时间性变化对结籽率的影响进行了研究。伊犁郁金香在居群不同时期的开放花密度、花蜜体积、花蜜有效性、访花频率、访问持续时间、传粉者数量及环境温度间均存在差异，其中温度及花蜜有效性是影响传粉者活动的主要因素。主要传粉者意蜂在居群不同时期的寻食行为和访花频率及传粉者效率间存在显著差异；其中在始花期和盛花期的传粉者效率属于低移出高沉积类型，而在末花期的传粉者效率属于高移出低沉积类型。传粉者效率和花粉传递效率与花结籽率间存在负相关性；其中在不同时期开放花的花粉传递效率从始花期到末花期逐渐下降，而结籽率从始花期到末花期逐渐提高。总之，伊犁郁金香在末花期的高移出-低沉积的传粉者效率和低效率的花粉传递效率对雌性繁殖成功的影响比始花期和盛花期的高移出-低沉积传粉者效率和高效率的花粉传递更具优势。
4.在自然居群中，比较研究其花期依赖的传粉者活动对花粉限制和近交衰退的影响进行了控制性试验。在花期不同时期的传粉者种类及访花频率间存在差异，传粉者活动从始花期到末花期，随着环境温度的上升而逐渐旺盛。在花期不同时期的自交不亲和指数、果实和种子水平的花粉数量和质量限制间存在差异。从始花期到末花期，质量和数量水平的花粉限制随着传粉者访花频率的提高而逐渐下降。不同授粉处理花所产生的种子的萌发率及雌性适合度间也存在差异，其中增补花粉的花雌性适合度比自然授粉花高；近交衰退指数从始花期到末花期逐渐下降，其中扩散前近交衰退比扩散后近交衰退更具优势；花粉质量限制与近交衰退指数间呈正相关性，而花粉数量限制与近交衰退指数间存在负相关性。早春环境的传粉者活动不仅影响伊犁郁金香花粉数量和/或质量限制水平及近交衰退，而且还能影响其后代适合度。早期激活的近交衰退是伊犁郁金香为适应早春环境而进化出的繁殖对策。
5.本研究对花大小依赖的自然选择特点与传粉特性及交配系统的关系进行了野外和室内的控制性试验。伊犁郁金香花的雌雄蕊长度与花瓣长大小间存在正相关性，而雌雄蕊空间接近程度与花瓣大小间呈负相关性。花雌雄器官获得资源比例随着花的增大更偏向于雌性方向，存在着大小依赖的性分配特点。传粉者在不同大小花上的访问行为间存在差异；花报酬和传粉者访花频率与花大小间呈正相关性。在自然授粉和去雄自然授粉花柱头上沉积的花粉数及结籽率与花大小间存在正相关性，而自主自花授粉花柱头上沉积的花粉及结籽率与花大小间存在负相关性。自交不亲和及近交衰退指数与花大小间存在彼此间的权衡关系；种子来表现扩散前近交衰退与花大小间呈正相关性。因此，整个居群的交配方式随着植株上花的变大倾向异交方向进化。伊犁郁金香在花性分配、传粉特性、交配方式及后代适合度上表现出的花大小依赖的自然选择特点是该物种为适应传粉者极少早春环境的繁殖成功而进化出的自然选择对策。
Under the rigid and variable extreme environment of the Junggar Desert in early spring, the diversityspring-flowering plants is low, and the vegetation is patchily-distributed. Moreover, pollinator species arerare, and the quality and quantity of pollinators varies temporally and spatially. Tulipa iliensis is an earlyspring ephemeral perennial that is widely distributed in central Asia, including northern Xinjiang, China.This species has floral temporal closure, and there are significantly different flower sizes in a population.Density of flowering plant, and pollinatoractivity vary during the flowering season. The relationshipbetween mating system and pollination characters and floral syndrome and the contribution of temporalfloral closure to reproductive success were measured. Effect of temporal changes in pollinator activity andpollen transfer efficiency on seed set, effect of flowering time dependent pollinator activity on pollenlimitation and inbreeding depression, and influence of flower size dependent sex allocation to matingpatterns were studied via field and laboratory observations or controlled experiments using standardreproductive biology methods. Survival strategies of the species in the early spring environment wereexamined in a natural population. The main results are as follows:
1. This study determined the effect of floral syndrome and pollination characters on mating patternsand their adaptation to the spring environment of T.iliensis. Flowers opened and closed temporarilyaccording to variations in temperature, exhibited incomplete herkogamy and homogamy, and had temporalvariation in duration of time they were open. Petals, stamen, and pistil were fragrant, and nectar volume andpollinators visiting frequency gradually decreased with an increased in flower age. Pollen viability, stigmareceptivity, and nectar volume were significantly positively correlated with pollinator visiting frequency(P<0.01). The bee Apis mellifera facilitated selfing. Flowers produced fruits after self-pollination (autogamyand hand self pollination) and cross-pollination; thus, they have a mixed mating system. There were highlysignificant differences among seed germination percentages and female fitness of differently-treated flowers.This species exhibited pollen limitation due to low pollinator activity. It has an autonomously self pollinationfloral syndrome, and facilitated selfing by the visiting behavior of pollinators, which assures the reproductivesuccess of T.iliensis in the unpredictable harsh spring environment.
2. This study examined the contribution of temporal floral closure on reproductive success inlaboratory and field experiments. Floral closure could result in contact of the dehisced anthers with thestigma and auto-pollen deposition on the stigma, thus facilitating automatic self pollination. Further, flower closure could maintain the temperature inside the flower and thus provide a stable "microenvironment" thatwould enhance pollen germination and avoid a decline in stigma receptivity of T. iliensis. Also, flowerclosure before rainstorms could avoid flushing of pollen from anthers and maintain both pollen viability andstigma receptivity during exposure to rain. Flower closure could maintain the pollen clumping in anthersduring exposure to wind and enhance pollination efficiency or facilitate automatic selfing. Therefore,temporal flower closure is an important floral strategy for facilitating self-pollination and for extending thefunctional period of male and female organs, which significantly increases reproductive success in theunpredictable harsh spring environment.
3. This study explored the effects of temporal changes in pollinators and transfer pollen efficiency onseed set. The results show that the nectar volume and availability, temperature, visiting frequency, andpollinator characteristics change with different periods in the flowering season, with nectar availability andtemperature being major factors that influence pollinator activity. Flower visitation frequency and foragingbehavior of the major pollinator Apis mellifera differed significantly with flowering period. Pollinatorefficiency was reted as “low removal high deposition visitor” at early and peak period, and was “highremoval low deposition visitor” in the late period. Pollinator efficiency and pollen transfer efficiencywerenegatively significantly correlated with female reproduction and seed set gradually increased from the earlyto the late flowering periods. Low nectar availability and flowering density, high visiting frequency andpollinator density were important factors that affected female reproductive success and pollinator efficiency.Temporal change in pollinator activity may influence pollen transfer efficiency and seed set. Therefore, thehigh removal low deposition visitors and low pollen transfer efficiency were better in the late floweringperiod than in the early and peak flowering periods, which protects the reproductive success and thusincreases genetic diversity of T.iliensis in the unpredictable harsh spring environment.
4This study examined the effects of flowering-period dependent pollinator activity on pollen limitationand inbreeding depression. Pollinator density and visiting frequency were significantly different betweenflowering periods. Pollinators activity gradually increased from the early to the late period with the increaseof environmental temperature. There were significant differences between the self-incompatibility and pollenlimitation index for fruit and seed levels at different flowering periods. Thus, cumulative pollen limitationdepended more on pollen quality than on quantity; pollen limitation gradually increased from the early to thelate flowering period with increasing pollinator activity. Seed germination percentage and female fitness ofthree flower treatments differed significantly at different flowering periods, and it was higher forsupplementally pollinated flowers than for naturally pollinated flowers. Inbreeding depression decreasedfrom the early to the late flowering period, and early-acting inbreeding depression was higher thanlate-acting inbreeding depression. Inbreeding depression was significantly positively correlated with qualityof pollen limitation, but it was significantly negatively correlated with quantity of pollen limitation. Pollinator activity was the major factor for the reproductive success of T.iliensis, and it not only affectedpollen limitation and inbreeding depression of individuals but also offspring fitness of T.iliensis plants thatflower at different times. Early-acting inbreeding depression was an important natural selective factor for thedirection of evolution direction and environment adaptation of T.iliensis.
5. This study explored the effects of flower size dependent sex allocation on pollination, mating patterns,female reproduction and offspring fitness through laboratory and field experiments. Petal length (size) was18mm-48mm; flower size was significantly positively correlated with stamen and pistil length, but it wassignificantly negatively correlated with stamen-pistil spatial extent. The percentage of resources allocated tothe pistil was greater in large than in small flowers, and resource allocation for flowers of different sizes wasa trade-off. There were significant differences between pollinator visiting behavior and flower size. Thenumber of pollen grains deposited on the stigma and seed set for naturallly pollinated and for naturalllypollinated flowers that had been emasculated were significantly positively correlated with flower size.However, number of pollen grains deposited on the stigma and seed set via spontaneous autogamy withdifferent flower size. Selfing rate increased with flower size, and inbreeding depression was a trade-off.Flower size was the major factor changing mating patterns to out-crossing.
1.花部综合征和传粉特性对交配方式的影响及其在早春环境的适应进行了研究。该物种花具有温度变化导致的花暂时性闭合特性、花不完全雌雄异位、花内雌雄同熟及花开放持续时间的花龄依赖的变化等花部综合征；花瓣、雄蕊及雌蕊均能分泌香味。花蜜体积及传粉者访花频率随着花龄的增大而逐渐减少；其访花频率与花粉活力、柱头可授性及花蜜体积间均存在正相关关系。自花授粉与异花授粉花均结实，属于混合交配系统；存在自主自交及意蜂(Apis mellifera)在花上的运动模式引起的协助自交。不同处理的花所产生的种子萌发率间均存在显著差异；异花授粉种子的适合度比自然授粉和自花授粉种子高。花雌雄功能分离的时空特点、花报酬及花暂时性闭合等花部综合征有关的自主自交及传粉者行为导致的协助自交是该物种在长期适应传粉者限制的早春环境中，为保障繁殖成功而形成的环境适应对策。
2.花暂时性闭合对繁殖成功的贡献进行了野外和室内的控制性试验。在花完全开放和完全闭合期间的柱头和花药空间位置间存在差异；花暂时性闭合能促进自主自交。花暂时性闭合能保持花内微环境温度，提供较稳定的花微环境温度来影响花粉萌发率及柱头可授性。在下雨前伊犁郁金香通过花暂时性闭合来避免雨水对花药上花粉的冲刷以及对花粉活力和柱头可授性的影响。在大风前的花暂时性通过保护花粉聚集水平来影响传粉者的传粉效率或促进自主自交率。伊犁郁金香花的暂时性闭合是该物种在长期适应早春环境中，为了促进自主自交、延长雌雄器官的功能及提高繁殖成功而进化出的非花部结构机制。
3.在自然居群中，传粉者及花粉传递效率的时间性变化对结籽率的影响进行了研究。伊犁郁金香在居群不同时期的开放花密度、花蜜体积、花蜜有效性、访花频率、访问持续时间、传粉者数量及环境温度间均存在差异，其中温度及花蜜有效性是影响传粉者活动的主要因素。主要传粉者意蜂在居群不同时期的寻食行为和访花频率及传粉者效率间存在显著差异；其中在始花期和盛花期的传粉者效率属于低移出高沉积类型，而在末花期的传粉者效率属于高移出低沉积类型。传粉者效率和花粉传递效率与花结籽率间存在负相关性；其中在不同时期开放花的花粉传递效率从始花期到末花期逐渐下降，而结籽率从始花期到末花期逐渐提高。总之，伊犁郁金香在末花期的高移出-低沉积的传粉者效率和低效率的花粉传递效率对雌性繁殖成功的影响比始花期和盛花期的高移出-低沉积传粉者效率和高效率的花粉传递更具优势。
4.在自然居群中，比较研究其花期依赖的传粉者活动对花粉限制和近交衰退的影响进行了控制性试验。在花期不同时期的传粉者种类及访花频率间存在差异，传粉者活动从始花期到末花期，随着环境温度的上升而逐渐旺盛。在花期不同时期的自交不亲和指数、果实和种子水平的花粉数量和质量限制间存在差异。从始花期到末花期，质量和数量水平的花粉限制随着传粉者访花频率的提高而逐渐下降。不同授粉处理花所产生的种子的萌发率及雌性适合度间也存在差异，其中增补花粉的花雌性适合度比自然授粉花高；近交衰退指数从始花期到末花期逐渐下降，其中扩散前近交衰退比扩散后近交衰退更具优势；花粉质量限制与近交衰退指数间呈正相关性，而花粉数量限制与近交衰退指数间存在负相关性。早春环境的传粉者活动不仅影响伊犁郁金香花粉数量和/或质量限制水平及近交衰退，而且还能影响其后代适合度。早期激活的近交衰退是伊犁郁金香为适应早春环境而进化出的繁殖对策。
5.本研究对花大小依赖的自然选择特点与传粉特性及交配系统的关系进行了野外和室内的控制性试验。伊犁郁金香花的雌雄蕊长度与花瓣长大小间存在正相关性，而雌雄蕊空间接近程度与花瓣大小间呈负相关性。花雌雄器官获得资源比例随着花的增大更偏向于雌性方向，存在着大小依赖的性分配特点。传粉者在不同大小花上的访问行为间存在差异；花报酬和传粉者访花频率与花大小间呈正相关性。在自然授粉和去雄自然授粉花柱头上沉积的花粉数及结籽率与花大小间存在正相关性，而自主自花授粉花柱头上沉积的花粉及结籽率与花大小间存在负相关性。自交不亲和及近交衰退指数与花大小间存在彼此间的权衡关系；种子来表现扩散前近交衰退与花大小间呈正相关性。因此，整个居群的交配方式随着植株上花的变大倾向异交方向进化。伊犁郁金香在花性分配、传粉特性、交配方式及后代适合度上表现出的花大小依赖的自然选择特点是该物种为适应传粉者极少早春环境的繁殖成功而进化出的自然选择对策。
Under the rigid and variable extreme environment of the Junggar Desert in early spring, the diversityspring-flowering plants is low, and the vegetation is patchily-distributed. Moreover, pollinator species arerare, and the quality and quantity of pollinators varies temporally and spatially. Tulipa iliensis is an earlyspring ephemeral perennial that is widely distributed in central Asia, including northern Xinjiang, China.This species has floral temporal closure, and there are significantly different flower sizes in a population.Density of flowering plant, and pollinatoractivity vary during the flowering season. The relationshipbetween mating system and pollination characters and floral syndrome and the contribution of temporalfloral closure to reproductive success were measured. Effect of temporal changes in pollinator activity andpollen transfer efficiency on seed set, effect of flowering time dependent pollinator activity on pollenlimitation and inbreeding depression, and influence of flower size dependent sex allocation to matingpatterns were studied via field and laboratory observations or controlled experiments using standardreproductive biology methods. Survival strategies of the species in the early spring environment wereexamined in a natural population. The main results are as follows:
1. This study determined the effect of floral syndrome and pollination characters on mating patternsand their adaptation to the spring environment of T.iliensis. Flowers opened and closed temporarilyaccording to variations in temperature, exhibited incomplete herkogamy and homogamy, and had temporalvariation in duration of time they were open. Petals, stamen, and pistil were fragrant, and nectar volume andpollinators visiting frequency gradually decreased with an increased in flower age. Pollen viability, stigmareceptivity, and nectar volume were significantly positively correlated with pollinator visiting frequency(P<0.01). The bee Apis mellifera facilitated selfing. Flowers produced fruits after self-pollination (autogamyand hand self pollination) and cross-pollination; thus, they have a mixed mating system. There were highlysignificant differences among seed germination percentages and female fitness of differently-treated flowers.This species exhibited pollen limitation due to low pollinator activity. It has an autonomously self pollinationfloral syndrome, and facilitated selfing by the visiting behavior of pollinators, which assures the reproductivesuccess of T.iliensis in the unpredictable harsh spring environment.
2. This study examined the contribution of temporal floral closure on reproductive success inlaboratory and field experiments. Floral closure could result in contact of the dehisced anthers with thestigma and auto-pollen deposition on the stigma, thus facilitating automatic self pollination. Further, flower closure could maintain the temperature inside the flower and thus provide a stable "microenvironment" thatwould enhance pollen germination and avoid a decline in stigma receptivity of T. iliensis. Also, flowerclosure before rainstorms could avoid flushing of pollen from anthers and maintain both pollen viability andstigma receptivity during exposure to rain. Flower closure could maintain the pollen clumping in anthersduring exposure to wind and enhance pollination efficiency or facilitate automatic selfing. Therefore,temporal flower closure is an important floral strategy for facilitating self-pollination and for extending thefunctional period of male and female organs, which significantly increases reproductive success in theunpredictable harsh spring environment.
3. This study explored the effects of temporal changes in pollinators and transfer pollen efficiency onseed set. The results show that the nectar volume and availability, temperature, visiting frequency, andpollinator characteristics change with different periods in the flowering season, with nectar availability andtemperature being major factors that influence pollinator activity. Flower visitation frequency and foragingbehavior of the major pollinator Apis mellifera differed significantly with flowering period. Pollinatorefficiency was reted as “low removal high deposition visitor” at early and peak period, and was “highremoval low deposition visitor” in the late period. Pollinator efficiency and pollen transfer efficiencywerenegatively significantly correlated with female reproduction and seed set gradually increased from the earlyto the late flowering periods. Low nectar availability and flowering density, high visiting frequency andpollinator density were important factors that affected female reproductive success and pollinator efficiency.Temporal change in pollinator activity may influence pollen transfer efficiency and seed set. Therefore, thehigh removal low deposition visitors and low pollen transfer efficiency were better in the late floweringperiod than in the early and peak flowering periods, which protects the reproductive success and thusincreases genetic diversity of T.iliensis in the unpredictable harsh spring environment.
4This study examined the effects of flowering-period dependent pollinator activity on pollen limitationand inbreeding depression. Pollinator density and visiting frequency were significantly different betweenflowering periods. Pollinators activity gradually increased from the early to the late period with the increaseof environmental temperature. There were significant differences between the self-incompatibility and pollenlimitation index for fruit and seed levels at different flowering periods. Thus, cumulative pollen limitationdepended more on pollen quality than on quantity; pollen limitation gradually increased from the early to thelate flowering period with increasing pollinator activity. Seed germination percentage and female fitness ofthree flower treatments differed significantly at different flowering periods, and it was higher forsupplementally pollinated flowers than for naturally pollinated flowers. Inbreeding depression decreasedfrom the early to the late flowering period, and early-acting inbreeding depression was higher thanlate-acting inbreeding depression. Inbreeding depression was significantly positively correlated with qualityof pollen limitation, but it was significantly negatively correlated with quantity of pollen limitation. Pollinator activity was the major factor for the reproductive success of T.iliensis, and it not only affectedpollen limitation and inbreeding depression of individuals but also offspring fitness of T.iliensis plants thatflower at different times. Early-acting inbreeding depression was an important natural selective factor for thedirection of evolution direction and environment adaptation of T.iliensis.
5. This study explored the effects of flower size dependent sex allocation on pollination, mating patterns,female reproduction and offspring fitness through laboratory and field experiments. Petal length (size) was18mm-48mm; flower size was significantly positively correlated with stamen and pistil length, but it wassignificantly negatively correlated with stamen-pistil spatial extent. The percentage of resources allocated tothe pistil was greater in large than in small flowers, and resource allocation for flowers of different sizes wasa trade-off. There were significant differences between pollinator visiting behavior and flower size. Thenumber of pollen grains deposited on the stigma and seed set for naturallly pollinated and for naturalllypollinated flowers that had been emasculated were significantly positively correlated with flower size.However, number of pollen grains deposited on the stigma and seed set via spontaneous autogamy withdifferent flower size. Selfing rate increased with flower size, and inbreeding depression was a trade-off.Flower size was the major factor changing mating patterns to out-crossing.
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