天仙子的花部特征及延迟自交机制

通过实验观察和野外控制授粉等方法,对天仙子的自交亲和能力和自交传粉机制进行了研究。结果显示:(1)天仙子的花不完全雌性先熟,开花进程中雄蕊伸长是维持其混合交配系统的重要花部特征。(2)野外控制授粉实验证明天仙子的自交亲和性高,具有主动自交能力,主动自交发生的时间集中在晚花期,与花药和柱头接触的时间吻合。(3)在花粉活力和柱头可授期内,花药与柱头的接触能够发生自花授粉,这种依赖花药与柱头接触发生的自花授粉机制属于典型的延迟自交类型。研究表明,天仙子不仅在花结构上首先选择了适于异交的特征,而且还利用雄蕊伸长成功实现自花授粉,通过异交与延迟自交混合的交配系统为该物种提供了灵活的遗传与繁殖保障。     

气候条件对海滨锦葵(Kosteletzkya virginica) 延迟自花传粉的影响

延迟自交被认为是一种生殖适应性,因为它保证了传粉者稀少情况下的种子生产,但当传粉者丰富时,又允许异交占优势。海滨锦葵花在传粉者不足或缺乏时能通过主动的柱头裂片弯曲运动成功实现延迟自花传粉。2002-2005年对江苏盐城滩涂海滨锦葵栽培群体花中发生延迟自花传粉花所占的百分比进行了观测,并于2005年对花内不同停止类型的柱头裂片比进行了调查,结果表明,2002-2005年间发生延迟自花传粉花所占百分比分别为13.50%±1.62%、14.39%±1.50%、14.05%±1.82%和13.47%±1.51%,不同年份间并无显著差异（F（3,189）=6.128,P=0.085）。发生延迟自花传粉花所占百分比与气候条件有密切关系,不利天气（多云/雨）（22.11%±0.94%）明显高于有利天气（晴天）（5.43%±0.43%）（F（1,189）=8009.780,P〈0.001）。花内通过花部运动实现自花传粉而停止运动的柱头裂片比在有利天气（晴天）下（18.64%±1.61%）明显低于不利天气（多云/雨天）（70.58%±2.06%）（U=0.000,W=465.000,P〈0.001）。海滨锦葵这种直接响应于影响传粉者环境之气候条件的延迟自花传粉策略为从个体或花水平上验证繁殖保障假说提供了可能。     

入侵植物美洲商陆的传粉生态学特性

从开花物候、花朵数量性状、花粉活力、花粉组织化学、柱头可授性、花粉在柱头上的萌发状况、杂交指数、花粉胚珠比、传粉媒介以及繁育系统等方面,研究了外来入侵植物美洲商陆(Phytolacca americana)的传粉生态学特性,为揭示该物种成功入侵的机理提供依据.结果表明:美洲商陆种群花期可达45~70 d,单花花期为2~3d,可进一步分为杯状花期、花瓣平展期、花瓣下垂期、花瓣反卷期;花冠直径在下垂期最大且不超过1 cm,花药与柱头间距离在1 ~2.5 mm,有利于自交的发生;花瓣平展期与花瓣下垂期花粉活力差异显著(P<0.05),杯状花期与花瓣平展期花粉活力差异不显著(P>0.05),花瓣平展期花粉活力可达85%;雌蕊柱头可授性在花瓣平展期最强;杯状花期合生柱头开裂,花瓣平展期柱头9~ 10裂,接受花粉的表面积增大;花药散粉时间与合生柱头开裂时间相吻合,有助于柱头接受花粉;扫描电镜下观察到花粉在花瓣平展期的柱头上已萌发;每朵花可提供1667~3222粒花粉,胚珠9~10个,花粉胚珠比(P/O)为340.88±34.99,表现为兼性自交;美洲商陆杂交指数(OCI)为3,花粉粒属于含脂型,主要传粉昆虫有蜜蜂、胡蜂、食蚜蝇、细腰蜂.综上所述,美洲商陆自交亲和,花粉活力高、柱头可授性强,且高花粉活力与强柱头可授性出现在相同的时期,增加了花粉落置柱头及在柱头上萌发的几率;虫媒花,访花者行为与花朵相适应,有利于提高坐果率,这样的繁殖机制与传粉特点是美洲商陆成功入侵的主要因素之一.     

山羊臭虎耳草的延迟自动自花授粉机制

通过对山羊臭虎耳草6个花形态及其发育阶段、授粉过程的连续观察和对花粉活性、柱头可授性的连续检测,报道了一种新的延迟自动自花授粉机制,即山羊臭虎耳草花丝运动引导的延迟自动自花授粉。利用电子显微镜研究了花粉囊的破裂方式和柱头结构,同时利用苏丹Ⅲ研究了花粉的组织化学性质。结果显示:(1)第四生长阶段,当花丝引导大部分花粉囊运动至柱头上方时,花粉和柱头的可授性才均达到峰值[可授性分别为(94±6)%和(96±13)%],这种运动消除了第三生长阶段花粉活性较高[(90±8)%]而柱头可授性较低[(6±17)%]的异花授粉特征,即消除了花粉与柱头的时空隔离;自动自花授粉发生于异花授粉特征消除之后,因此山羊臭虎耳草具有典型的延迟自动自交行为。(2)第四发育阶段柱头空腔的大量柱状突起和花粉鞘的油脂性特征,增加了花粉和柱头的接触表面,并且使花粉更好地互相堆积粘附在柱头上,提高授粉效率。(3)对山羊臭虎耳草的繁育系统研究表明,强制自花授粉和人工自花授粉产种量无明显差异,2007年和2008年的自交系数高达0.92和0.91,说明山羊臭虎耳草具有很好的自交能力并且不存在花粉数量限制;人工异花授粉的产种率显著高于人工自花授粉,说明自花花粉的质量显著低于异花花粉,虎耳草延迟自动自花授粉存在生殖衰退,生殖保障系数2007年为0.92,2008年为0.91,表明在极端环境下自动自花授粉为该植物提供了繁殖保障,但极端环境下,没有观察到有效的授粉昆虫。研究表明,作为高山冰缘植物的山羊臭虎耳草,保留了开花植物异花授粉的特征,在极端环境下利用延迟自动自花授粉机制完成授粉,保障生殖。     

Differences in floral traits and flower visitation rates in mating systems in Prunella vulgaris (Lamiaceae)

夏枯草交配系统对花特征和访花频率差异的影响 植物花特征和传粉者的访问次数与交配系统类型密切相关。唇形科植物夏枯草(Prunella vulgaris)存 在两种植株类型,分别为柱头伸出花冠和柱头在花冠内部的植株,而且两种植株的比例在不同种群中存在差异。本研究选择柱头伸出花冠外植株占绝大多数、柱头伸出花冠外植株占多数和柱头在花冠内部植株占多数的3个种群,通过比较每个种群中两种植株类型的开花物候、花形态特征、昆虫访问频率、自交能力、传粉者对结实的贡献以及近交衰退的水平,以检验花特征和传粉者访问次数与交配系统类型的关系。研究结果表明,与柱头在花冠内部的植株相比,柱头伸出花冠外的植株具有更大和更多的花,产生更多的花粉和花蜜,具有更高的访花频率,并主要通过异交产生种子。在种群水平,柱头伸出花冠外的植株占多数种群的访花频率显著高于柱头在花冠内部植株占多数的访花频率。柱头在花冠内部的植株比柱头伸出花冠外的植株具有更强的自动自交能力,在传粉者缺乏时为其提供了繁殖保障,但繁殖保障和异交率在不同种群中差异不显著,表明较低的昆虫访问能够满足夏枯草的授粉需求以产生种子,这可能与夏枯草较少的胚珠数量(每朵花仅有4个胚珠)有关。柱头在花冠内部植株的近交衰退水平低于柱头伸出花冠外植株的近交衰退水平,但两种植株类型的近交衰退水平均低于0.5,说明近交衰退不足以阻止该物种中自交的进化。综上所述,柱头在花冠内部的植株能够通过自交为夏枯草提供繁殖保障,而柱头伸出花冠外的植株能够利用昆虫传粉确保异交,表明混合交配系统在该物种中是一个稳定的状态。     

早春不稳定传粉环境中新疆郁金香混合交配系统中的自交策略

【目的】在植物的所有生活史特征中,交配格局可能是对宏观进化影响最大的因素。在不确定的传粉环境中,两性花植物常常具有潜在的自交能力,鉴于自交的交配代价,两种交配方式如何权衡,一直是深入理解交配系统演化的关键问题。【方法】为了探讨早春不稳定传粉环境中植物的自交策略,以早春短命植物新疆郁金香为研究对象,通过野外观测及人工控制实验对自然居群的开花习性、传粉者类群、散粉动态、自花粉传递模式和交配系统等进行研究。【结果】（1）新疆郁金香自然种群4月上旬或中旬开花,单花期5-6 d,白天开放晚上闭合,花粉的释放从外轮开始,由下到上呈拉链式次序呈现。（2）传粉者主要为蜂类和食蚜蝇,访花频率普遍较低,且在年份间存在较大差异,但结实率普遍较高。（3）控制授粉实验表明居群为异交为主,部分自交亲和的混合性交配系统类型。傍晚花闭合时雄蕊的自主运动促进了柱头的自花授粉,这一传粉模式促进了竞自交的发生,但大量自花粉的落置发生在开花后的第4 d,占自花粉总落置量的50.22%,为一种延迟自交的机制。【结论】在传粉受限的情况下,新疆郁金香的竞自交和延迟自交促进了柱头的花粉落置,这种集异交、竞自交和延迟自交为一体的交配策略灵活地应对了早春不稳定的传粉环境,是对早春低温度条件下不确定传粉服务的一种适应,也是早春短命植物的一种繁殖保证对策。     

红豆蔻专性异交主导下的主动自交机制（英文）

主动自交一直以来都被认为是植物在面对极端环境,例如传粉者稀缺的情况下所采取的繁殖保障机制,它不利的一面主要表现为近交衰退和花粉或胚珠贴现。在姜科植物部分属,如山姜属和豆蔻属当中,存在着一种被称作花柱卷曲(flexistyly)的特殊机制。过去的研究认为这种机制的意义在于避免自交和性别干扰,但是根据我们的野外观察,如果下垂型花朵的花粉在其花柱开始向下弯曲时仍然留在花药之上,柱头就很有可能接触到花粉从而完成主动自交。为此我们探索了红豆蔻(Alpinia galanga)的繁殖系统,研究结果表明:(1)红豆蔻自交亲和,并存在一定的近交衰退;(2)上举型为异交,较之下垂型投入了更多的资源;(3)上举型的花粉胚珠比显著低于下垂型,由于红豆蔻的胚珠数是固定的六枚,故两种表型花粉胚珠比的差异反映了其花粉数目的差异。这可能与上举型仅仅为下垂型提供花粉,而下垂型除为上举型提供花粉外,还需要有足够的花粉以备自交所需。因此,红豆蔻当中存在主动自交机制,并且其花柱卷曲机制在避免不必要的自交和性别干扰的同时,也为作为繁殖保障所必需的延迟主动自交创造了有利条件。同时通过控制主动自交发生的时间达到避免三种功能之间的相互冲突。这种只在姜科植物当中存在的特殊机制充分展示了其进化过程中对不利环境的适应。     

短命植物甘新念珠芥(Neotorularia korolkovii)和宽翅菘蓝(Isatis violascens)的物候与性表达特征

甘新念珠芥（Neotorularia korolkovii）和宽翅菘蓝（lsatis violascens）是两种广泛分布于新疆准噶尔荒漠的十字花科早春短命植物。对其物候与性表达特征研究结果表明：（1）二者种子均在3月下旬萌发,5月下旬和6月上旬果实完全成熟,生活周期分别为62d和71d,其中营养生长期约占1／3、生殖生长期约占2／3。（2）花均为两性,四强雄蕊,单花开放时间较短,长雄蕊的花药开裂早于短雄蕊,但短雄蕊花药中的花粉数较长雄蕊的多。（3）长、短雄蕊花药中花粉可育率均达90％以上,花粉活力的变化曲线基本相同,柱头可授期分别为9．Oh和4．5h,花粉寿命及柱头可授期均短于单花花期,花粉活力最高的时期正是其柱头的最佳授粉期。柱头可授期与花粉活力及寿命在时间上的高度吻合,增加了在不可预测环境中授粉的机会与效率,从而保障生殖成功。（4）二者的P／O（花粉／胚珠）值分别为108．07±17．17、992.10±272．16,属于自交繁育系统;对其传粉过程、套袋实验及荧光显微观察的结果也证明它们主要以白花传粉的方式来进行交配。（5）两种植物从物候、花部性表达以及交配式样上均存在与其严酷荒漠环境相适应的特点,是长期适应准噶尔荒漠特殊生态环境的结果,对于保证繁殖成功并扩大种群具有重要的意义。     

珍稀濒危植物陕西羽叶报春的开花及传粉生物学研究

陕西羽叶报春为中国珍稀濒危植物,研究其开花物候、传粉者及访花行为、花粉活力、柱头可授性、繁育系统,以明确陕西羽叶报春的传粉特性及制濒机制。以勉县陕西羽叶报春野生居群为研究对象,通过野外观察和人工授粉实验首次对陕西羽叶报春的传粉生物学进行研究。结果表明:(1)陕西羽叶报春种群花期为2-4月中旬,单花花期一般为14~17 d;单花开花进程中花部形态有6个明显发育变化:全部被萼片包裹、花苞伸出并为深粉色、花朵打开为淡粉色、花冠为白色、花朵枯萎、花冠脱落花萼宿存。(2)陕西羽叶报春为异花授粉植物,蜜蜂属为其主要传粉者。(3)柱头在花粉未开裂前就具有可授性,可授性逐渐增强;花药在花瓣打开后的第3天才开始开裂、散粉,在第4天活力最高,为98.18%,此后逐渐减弱。(4)繁育系统检测显示,长、短柱型花的花粉/胚珠比分别为902.26和831.48,杂交指数为4;套袋实验显示,陕西羽叶报春异型交配可育,同型交配不可育。由此表明,陕西羽叶报春的繁育系统为异交型,自交不亲和,需要传粉者。综上所述,陕西羽叶报春传粉者种类少、自交不亲和、胚珠败育现象可能是导致陕西羽叶报春濒危的重要原因。     

气候条件对海滨锦葵（Kosteletzkya virginica）延迟自花传粉的影响

延迟自交被认为是一种生殖适应性,因为它保证了传粉者稀少情况下的种子生产,但当传粉者丰富时,又允许异交占优势。海滨锦葵花在传粉者不足或缺乏时能通过主动的柱头裂片弯曲运动成功实现延迟自花传粉。2002~2005年对江苏盐城滩涂海滨锦葵栽培群体花中发生延迟自花传粉花所占的百分比进行了观测,并于2005年对花内不同停止类型的柱头裂片比进行了调查,结果表明,2002 ~ 2005年间发生延迟自花传粉花所占百分比分别为13.50%±1.62%、14.39%±1.50%、1405%±1.82%和13.47%±1.51%,不同年份间并无显著差异 (F _(3, 189) = 6.128, P = 0.085 )。发生延迟自花传粉花所占百分比与气候条件有密切关系,不利天气(多云/雨) (22.11%±0.94%) 明显高于有利天气(晴天) (5.43%±0.43%) (F_(1, 189) = 8009.780, P < 0.001)。花内通过花部运动实现自花传粉而停止运动的柱头裂片比在有利天气(晴天)下(18.64%±1.61%)明显低于不利天气(多云/雨天) (70.58%±2.06%) (U = 0.000, W = 465.000, P <0.001)。海滨锦葵这种直接响应于影响传粉者环境之气候条件的延迟自花传粉策略为从个体或花水平上验证繁殖保障假说提供了可能。     

Movement herkogamy in Kosteletzkya virginica: Effect on reproductive success and contribution to pollen receipt and reproductive assurance

Herkogamy may act as a mechanism to reduce interference between the reproductive functions of female and male organs, but too great a distance between stigmas and anthers may also decrease the outcross pollen transfer and male and female fitness. Our goal was to quantify the effect of movement herkogamy in Kosteletzkya virginica on reproductive success and its contribution to pollen receipt and reproductive assurance. Field investigations and hand-pollination experiments were performed to test: (1) the relationship between herkogamy before curvature of un-pollinated lobes and reproductive success under pollinator exclusion; (2) the contribution of movement herkogamy to pollen receipt; and (3) the relationship between pollen load and seed set. We noted that variation in herkogamy before curvature of un-pollinated lobes did not affect the outcome of autogamous self-pollination measured by fruit set and mean number of seeds per fruit. Under natural conditions, 39% of pollen grains deposited per flower were from autogamous self-pollination via movement herkogamy resulting in contact between anthers and stigmas. The relationship between the amount of pollen grains and the number of seeds per capsule was highly significant. Flowers producing four or five seeds received more pollen grains than those producing only one seed. Our data provides experimental evidence that movement herkogamy, which decreases the separation between anthers and stigmas and results in delayed self-pollination, contributes to pollen receipt and further enhances seed set when pollinators are scarce or absent.     

Movement herkogamy in Kosteletzkya virginica: Effect on reproductive success and contribution to pollen receipt and reproductive assurance

Herkogamy may act as a mechanism to reduce interference between the reproductive functions of female and male organs, but too great a distance between stigmas and anthers may also decrease the outcross pollen transfer and male and female fitness. Our goal was to quantify the effect of movement herkogamy in Kosteletzkya virginica on reproductive success and its contribution to pollen receipt and reproductive assurance. Field investigations and hand-pollination experiments were performed to test: (1) the relationship between herkogamy before curvature of un-pollinated lobes and reproductive success under pollinator exclusion; (2) the contribution of movement herkogamy to pollen receipt; and (3) the relationship between pollen load and seed set. We noted that variation in herkogamy before curvature of un-pollinated lobes did not affect the outcome of autogamous self-pollination measured by fruit set and mean number of seeds per fruit. Under natural conditions, 39% of pollen grains deposited per flower were from autogamous self-pollination via movement herkogamy resulting in contact between anthers and stigmas. The relationship between the amount of pollen grains and the number of seeds per capsule was highly significant. Flowers producing four or five seeds received more pollen grains than those producing only one seed. Our data provides experimental evidence that movement herkogamy, which decreases the separation between anthers and stigmas and results in delayed self-pollination, contributes to pollen receipt and further enhances seed set when pollinators are scarce or absent.     

Delayed Selfing in an Alpine Biennial Gentianopsis paludosa (Gentianaceae) in the Qinghai-Tibetan Plateau

Delayed selfing could provide ovules with an opportunity to be fertilized as a means of "pollination assurance" before the flowers wilt. It could, thus, be regarded as an adaptation to unpredictable pollinator environments. Within the alpine biennial Gentianopsis paludosa, the showy flowers and herkogamy at the early stage of a flower's life cycle may favor outcrossing. As the flower ages, anthers contact the central stigma due to the elongation of all filaments, resulting in autonomous selfing. Flower visitors are extremely rare in a high altitude population; and examination of the mating system indicates that G. paludosa is self-pollinated under natural conditions in this population. While at the lower altitude, the bumblebee visitation rate is relatively high but possibly unreliable. Stigma receptivity is the highest on the third day of anthesis, and decreases thereafter. Pollen viability is the highest when flowers open, and gradually decreases later. Self-pollination of G. paludosa occurs at the late stage of a flower's lifecycle when stigma receptivity and pollen viability have both decreased, suggesting delayed selfing and assurance of seed production. This delayed selfing could assure seed production under the constraints of pollinator scarcity, but ensure outcrossing when pollinators were available. Such a flexible pollination mechanism is highly adaptive in the alpine environment of the Qinghai-Tibetan Plateau.     

高山植物扁蕾的延迟自交机制

扁蕾(Gentianopsis barbata)具有鲜艳的花和显著的腺体,并且花开放的前5 d柱头和花药始终处于不同的位置(雌雄异位),这些花综合征表明该植物应为异花传粉。为检验这一假设,我们对青藏高原植物扁蕾的海北站种群进行了3年的传粉生物学研究实验。与花综合征所表明的繁育系统相反,两年的野外观察发现昆虫的访花频率十分低,不去雄并隔离昆虫处理也能产生大量种子,说明这一种群的繁殖主要是依赖于自花传粉。尽管利用种子结实评价的柱头可授性从花开放4 d后开始下降,但随着花的发育进程,雄蕊的伸长能使得花药与柱头完全接触。实验也证明,柱头可授性和花粉活力都超过5 d,说明花药和柱头的接触能够发生自花授粉。扁蕾的这种自花传粉机制应属于典型的延迟自交类型。自花授粉发生在单花花期快要结束前,自交之前仍然保持异交传粉机制,这种延迟自交避免了自交与异交竞争造成的花粉或者种子折损,并为扁蕾在青藏高原极端环境下由于访花昆虫缺乏造成的异交失败提供了繁殖保障。     

Context-dependent style curvature in Kosteletzkya virginica (Malvaceae) offers reproductive assurance under unpredictable pollinator environments

Delayed selfing is a reproductive strategy that can increase mating opportunities and provide reproductive assurance in the absence of pollinators. When external pollination fails to occur in the herbaceous perennial, Kosteletzkya virginica, the styles curve downward to bring anthers into contact with the stigmal lobes, resulting in delayed selfing. Our goal was to evaluate experimentally the effects of weather conditions on pollinator activity, style curvature, and plant fitness. We discovered that adverse weather conditions significantly reduced pollinator visitation rates, and increased the incidence of style curvature, which resulted in delayed selfing. Furthermore, elimination of delayed selfing by experimental manipulation reduced fruit and seed set compared to non-emasculated flowers, even under adverse weather conditions. Offspring produced through selfing were similar to outcrossed progeny in the majority of fitness traits measured, and inbreeding depression was limited. Our study demonstrates that context-dependent style curvature promotes delayed selfing and provides reproductive assurance in the absence of pollinators, with minimal loss of progeny performance through inbreeding depression.     

Delayed autonomous self-pollination in the colonizer Crotalaria micans (Fabaceae: Papilionoideae): structural and functional aspects

We investigated the reproductive biology of Crotalaria micans, a colonizing species that occurs in disturbed sites. The flowers have two whorls of stamens with differential growth and dimorphic anthers. The inner anthers serve two functions: 1) to cooperate with the stylar brush in the pollen presentation, and 2) to push the remaining pollen within the keel onto the stigma at the end of the receptivity period. Flowers pass through three phases: a) male phase, when outer anthers release their pollen, but stigmas are not receptive; b) female phase, when stigmas are receptive, but still separated from flowers' own pollen, and c) an autogamous phase, when the round anthers grow towards the stigma, leading to delayed autonomous self-pollination. Because C. micans is completely self-compatible, delayed self-pollination and geitonogamy result in approximately 76% of seeds being self-fertilized.     

Does self-pollination provide reproductive Assurance in Aquilegia Canadensis (Ranunculaceae)?

The ability to produce seeds when pollinators or potential mates are scarce is thought to be one of the main advantages of self-fertilization in flowering plants. However, whether autonomous selfing increases seed set in natural populations has seldom been tested, and even fewer studies have evaluated the advantage of selfing across a gradient of pollen availability. This study examines the fertility consequences of autonomous selfing in Aquilegia canadensis (Ranunculaceae), a short-lived, spring-flowering perennial typically found in small, patchy populations on rock outcrops. We used a pollinator exclusion experiment to confirm reports that A. canadensis has a well-developed capacity for autonomous selfing resulting from incomplete protogyny and close proximity of stigmas and anthers during dehiscence. Flowers excluded from pollinators set 87% as many seeds per carpel (X +/- 1 SE = 7.1 +/- 1.4 seeds) as hand-pollinated flowers (8.1 +/- 1.3 seeds), and seed production in unpollinated flowers correlated negatively with the distance between stigmas and anthers (r = -0.46). Autonomous selfing could be potentially valuable in providing reproductive assurance because only 2.7 +/- 0.5 pollen grains were deposited on each stigma before anther dehiscence, compared to 134.1 +/- 17.9 pollen grains by the end of anther dehiscence. However, prevention of autonomous selfing by anther removal before dehiscence did not decrease seed set, even for plants at low plant densities where outcross pollen may have been in short supply. Emasculated flowers set as many seeds per carpel (9.3 +/- 0.9) as intact flowers (8.4 +/- 1.1), indicating that sufficient cross pollen is deposited for full seed set. These results do not support the hypothesis that autonomous selfing by A. canadensis has been selected because it provides reproductive assurance.     

The mechanism of delayed selfing in Collinsia verna (Scrophulariaceae)

Collinsia verna, blue-eyed Mary, has floral attributes of an outcrossing species, yet most flowers readily self-pollinate under greenhouse conditions. Here we describe the mechanism of self-pollination in C. verna via changes in relative positions of the stigma and anthers and late timing of receptivity, resulting in delayed selfing. Each flower contains four anthers that dehisce sequentially over ∼1 wk. Pollen that is not collected by pollinators accumulates in the keel petal and retains high viability (>80% pollen germination) up to the time of corolla abscission. The stigmatic surface does not become receptive until after the third anther dehisces. This overlap in the sexual phases is concurrent with a change in herkogamy during floral development. In most flowers (70%), the stigma has moved to the front of the keel and is positioned near the anthers when the third anther dehisces. Under field conditions, fruiting success of plants within pollinator exclosures was ∼75% of the fruiting success in open-pollinated plants (33% fruiting success via autogamy vs. 44% fruiting success, respectively). Collinsia verna plants in pollinator exclosures exhibit variation in autogamy rates within natural populations (range 0–80%). In addition, only half of naturally pollinated, receptive flowers examined had pollen tubes growing in their styles. In contrast, shortly after corolla abscission, nearly all flowers examined (96%) had pollen tubes in their styles. Thus we find that in C. verna, autogamy occurs late in floral development, which has the potential to provide substantial reproductive assurance, and that individuals vary in their ability to set fruit through this mechanism. We suggest that delayed selfing mechanisms may be overlooked in other species and that variable pollinator availability may play a significant role in the maintenance of mixed mating in species with delayed selfing, such as C. verna.     

Variation in the timing of autonomous selfing among populations that differ in flower size, time to reproductive maturity, and climate

? Premise of the study: Early reproductive maturity is common in dry and ephemeral habitats and often associated with smaller flowers with increased potential for within-flower (autonomous) self-pollination. We investigated whether populations from locations that differ in moisture availability, known to vary for whole-plant development rate, also varied in the timing of autonomous selfing. This timing is of interest because the modes of selfing (prior, competing, and delayed) have different fitness consequences. ? Methods: We measured timing of anther dehiscence, stigma receptivity, and herkogamy under pollinator-free conditions for plants from three populations of Collinsia parviflora that differed in annual precipitation, flower size, and time to sexual maturity. Using a manipulative experiment, we determined potential seed production via prior, competing, and delayed autonomous selfing for each population. ? Key results: Stigma receptivity, anther dehiscence, and selfing ability covaried with whole-plant development and climate. Plants from the driest site, which reached sexual maturity earliest, had receptive stigmas and dehiscent anthers in bud. Most seeds were produced via prior selfing. The population from the wettest site with slowest development was not receptive until after flowers opened. Although competing selfing was possible, all selfing was delayed. The intermediate population was between these extremes, with significant contributions from both competing and delayed selfing. ? Conclusions: Our results demonstrate that within-species variation in the timing of selfing occurs and is related to both environmental conditions and whole-plant development rates. We suggest that, if these results can be generalized to other species, mating systems may evolve in response to ongoing climatic change.     

繁殖保障和延迟自交的研究进展

尽管植物在进化过程中面临不利自花授粉的选择,但许多植物仍维持混合的授粉机制。繁殖保障假说是解释自交进化的最重要因子之一,一直是植物生殖生态学和进化生物学关注的焦点之一。概述了近年来的主要研究热点及其进展,包括自交进化的遗传和生态机制及理论模型探讨、繁殖保障假说的提出、验证自交能否提供繁殖保障的例证、延迟自交的类型及延迟自交能否提供繁殖保障的例证等方面。介绍了我国在繁殖保障和延迟自交方面研究的现状和不足之处,结合国际上研究繁殖保障假说的发展趋势已由单季节、单种群、单因子的研究阶段过渡到多季节、多种群、多因子（自交方式及其所占比例、花粉折损、种子折损、自交率和近交衰退）的综合研究阶段,及由传统的、经典的研究方法过渡到应用现代实验手段（如SSR、SNP等分子标记）和先进仪器设备的研究阶段,提出今后研究中应注意的问题。有必要借用多学科（植物学、生态学和分子生物学）的方法及手段进行不同物种的对比和综合细致的研究。