石榴花瓣和雄蕊对其传粉过程与繁殖成功的影响

植物-传粉者相互关系中植物形成多种多样的视觉(花色)和嗅觉(花气味)信号来影响传粉者的访花过程,促进传粉成功。石榴(Punicagranatum)花瓣红色而雄蕊黄色,这种花内不同结构的颜色差异对于石榴吸引传粉者可能有不同的作用。本文比较了石榴花各部位发出的视觉信号(颜色、大小)和嗅觉信号(气味及含量)、花蜜体积、不同处理下的昆虫访花频率及坐果率,以探讨石榴花各部位颜色在传粉过程中的作用。结果发现:新疆喀什地区石榴主要传粉者为意大利蜜蜂(Apis mellifera)和食蚜蝇(Syrphidae sp.),雄蕊黄色及其分泌挥发性化合物种类和相对含量是吸引传粉者昆虫的主要因素。去除花瓣处理组与其他3种处理组(对照、去雄蕊、去雄蕊去花瓣)比较,意大利蜜蜂的访花频率(P <0.05)及停留时间(P <0.05)均显著提高,石榴坐果率(82.33%±4.45%)也显著提高。上述结果表明,石榴黄颜色的雄蕊可能是吸引传粉者的主要结构,而红色的花瓣对其传粉成功可能有负面影响;植物花内不同结构颜色差异可能有助于在变化的环境下吸引不同的传粉者,促进繁殖成功。     

无花瓣油菜雄蕊心皮化突变体细胞学观察及基因表达分析

油菜是我国重要的油料作物,油菜花器官具有典型的十字花科特点,无花瓣油菜在花期不存在花冠层,这种特点有助于提高油菜产量,预防茵核病的传播。雄蕊心皮化是指花器官的雄蕊结构被具有类似于雌蕊结构的器官代替,这不仅造成了花器官结构的变化也导致了雄性不育。本文通过对无花瓣油菜雄蕊心皮化突变不育分离群体中的雄性可育株和不育株的比较研究,发现心皮化现象是由遗传因素引起的。细胞学观察发现,雄蕊心皮化在花器官发育的早期就已经产生,心皮化的雄蕊中着生类似于胚珠的结构,其顶端细胞的形态和排列方式也与雌蕊柱头相似。花发育相关基因的表达分析表明,B组基因彳丹在不育株3轮花器中的表达都比可育株低,特别是在第二轮花器官中这种差异最为突出。而A组基因AP1在不育株第二轮花器官中的表达量较可育株高。c组基因AGL8、SHPI、SHP2、NAP在不育株心皮化的第二轮花器官中表达都较可育株中高。     

光果甘草二体雄蕊的发育及其适应意义

甘草属(Glycyrrhiza)植物具“9 + 1”二体雄蕊, 其中9枚合生雄蕊的上部花丝分离, 分离的花丝在发育过程中存在由早期长、短交错的二组排列方式转变为后期以雄蕊管最长的1枚雄蕊为中心向两边渐次缩短的倒“V”形排列。为了解这种雄蕊发育动态、分化现象及发育成熟后的适应意义, 该文以光果甘草(G. glabra)为实验材料, 比较了雄蕊发育过程中的形态变化、成熟花粉的理化性质及在传粉中的作用。结果显示: 雄蕊发育早期长、短两组雄蕊在花药大小与形状上存在分化, 但后期伴随着花丝的快速生长与花粉的成熟、散出, 花药大小与形状趋于一致; 花粉组织化学成分及授粉成功率无差异, 但成熟花粉的数量和花粉活力存在差异; 去雄处理虽然使访花者在一天内的两个访花高峰期的访花频次降低, 但结实率高于自然对照, 说明以异交为主的花去除雄蕊后, 降低了雌、雄蕊间的功能干扰, 提高了传粉昆虫的授粉率; 发育早期长、短交错排列的二组雄蕊到成熟期时发生的倒“V”形排列的转变, 使不同数量与活性的花粉分布在花内不同空间, 最大化接触访花者, 实现了资源节约, 提高了雄蕊的雄性适合度, 即在有限的空间内用最节约的雄性资源投入、使传粉空间与传粉几率最大化的方式, 来提高雄性功能。     

栽培红花山玉兰的传粉生物学

研究了栽培红花山玉兰（Ｍａｇｎｏｌｉａｄｅｌａｖａｙｉ）的开花生物学和传粉生物学特性,并探讨了红花山玉兰只开花而不结实的原因以及提高其结实率的技术措施。红花山玉兰的花被片为９,呈３轮排列。外轮花被片先张开,而内２轮花被片仍紧密地包裹着雌雄蕊群,约２４ｈ后,内２轮花被片才张开。在外轮花被片张开而内２轮花被片未张开时,柱头就有授粉能力,但雄蕊尚未成熟。因内２轮花被片紧密地包裹着雌雄蕊群而阻碍了传粉者进入花内传粉。当内２轮花被片张开时,雄蕊成熟,花药裂开而散发花粉;柱头外露,但此时的柱头已变棕红色,完全失去了授粉能力。若外轮花被片刚张开时,去掉全部花被片,蜜蜂可成为有效传粉者,其结果率可达到５３．３％;若去掉全部花被片,并施以人工异花授粉,其结果率可高达１００％。研究结果表明,红花山玉兰是雌雄异熟的,且异花授粉是亲和;其开花生物学特性适合甲壳虫携带花粉进入花内传粉。红花山玉兰只开花而不结实的原因可能是其开花生物学特性所要求的携带花粉进入花内传粉的甲壳虫无法进入花内传粉或者缺乏。     

毛菍传粉生物学研究

为探讨毛菍(Melastoma sanguineum Sims.)传粉生物学特性,对海南霸王岭国家级自然保护区林缘、林内和林隙3个生境内毛菍的花期、花部形态、访花昆虫等进行观察,同时对其繁育系统相关指标进行了测定。结果表明：毛菍无花蜜分泌;花瓣对访花昆虫有吸引作用;访花昆虫主要有蜂类、食蚜蝇类和丽金龟类;主要传粉昆虫有木蜂、熊蜂,其中木蜂传粉活动最为活跃,为最有效传粉昆虫。毛菍具典型的异型雄蕊,长、短雄蕊在传粉过程中具有明确的功能分化,推测这是毛菍保证传粉成功的策略。毛菍为自交亲和的异交种,需要昆虫传粉;人工自交和异交均具有较高的坐果率;不存在无融合生殖、主动自花授粉和滞后自交的生殖保障现象;其繁育系统是兼性自交。昼夜温差较小和相对湿度较高的林内和林隙生境有利于延长群体花期;光照充足、温度相对较高的林缘生境,花朵发育较快,柱头可授性的时间稍早。花粉、传粉者和环境条件是制约毛菍野外传粉效率的主要因素。     

珍稀濒危植物小花木兰传粉生物学研究

基于定株观测,对小花木兰（Magnolia sieboldii）野生种群和人工栽培种的开花物候进行了研究。结果表明,小花木兰花期为5月中下旬至6月中下旬,花期20d左右,单花花期一般为5～7d,盛花期为6月1～5日;开花过程中花部表型变化明显,花粉在花开时不成熟,但雌蕊已经成熟,花开后雌蕊柄伸长,约20h后下柱头依然湿润,上柱头已萎缩变褐色,再过约40h下柱头也萎缩失去活力．此时花粉成熟,花药裂开;花的柱头接受到的花粉量由下往上依次减少。检测花粉活力、萌发率发现,小花木兰花粉在散发6d后基本失去活力和萌发能力。重力玻片法观测表明,小花木兰为虫媒传粉;花柱的授粉率不高,为65％;柱头上平均花粉量为3．5个;自然状态下结实率低,为13．5％,人工栽培为20．1％;外轮花被片全部去掉,结果率下降至11．3％;去掉雄蕊,结实率下降至10．7％;再辅以人工授粉,结果率上升至25．0％。小花木兰的访花昆虫种类较少,访花频率低,其中蜜蜂和一种蚜科（Aphididae）小昆虫对其传粉影响较大。同花期灯台附（Cornus controversa）等植物对其传粉产生昆虫竞争作用。环境因素如光照、温度、阴雨、海拔等均对其传粉产生一定影响。     

栽培红药山玉兰的传粉生物学

研究了栽培红山玉兰（Ｍａｇｎｏｌｉａ ｄｅｌａｖａｙｉ）的开花生物学和传粉生物特性,并探讨了红花山玉兰只开花而不结实的原因以及提高其结实率的技术措施。红花山玉兰的花被片为９,呈３轮排列。外轮花被片先张开,而内２轮花被片仍紧密地包裹着雌雄蕊群,约２４ｈ后,内２轮花被片才张开。在外轮花被片张开而以２轮花被片未张开时,柱头就有授粉能力,但雄蕊尚未成熟。因内２轮花被片紧密地包裹着雌雄蕊而阻碍了传粉者进入花     

单性木兰花部综合特征及其传粉适应性

对单性木兰花部综合特征和访花昆虫种类及访花行为进行了研究,结果表明,单性木兰花冠顶端呈半合拢状,花被片之间存在间隙,只有那些能够穿过这些间隙的昆虫才能到达花内为其采粉和授粉,对访花者的要求较为苛刻。于降雨前去掉雄花和雌花花被片,使雄花雄蕊群和雌花柱头充分暴露,另外选择雄花和雌花作对照,降雨后将花取回并测定雄花花粉生活力和雌花柱头上的花粉粒。结果表明,在雨天,单性木兰花冠顶端呈半合拢状的开放方式对雄花和雌花均起到了保护作用,这种花部结构的特化是对花期持续降雨的一种生态适应。从访花昆虫种类上看,单性木兰雄花的昆虫种类是雌花的两倍,显然雄花比雌花更能吸引昆虫。     

柴黄姜的传粉生物学特性

对产自湖北武当山的柴黄姜（Dioscorea nipponica subsp．rosthornii Prain et Burkill）的花部特征和访花昆虫种类及访花行为、访花频率和携粉量进行了研究,并对柴黄姜的传粉方式、结实和种子萌发状况进行了检测。结果表明,柴黄姜具有虫媒花的典型花部特征,共观测到访花昆虫4目34种,其中传粉昆虫6种,隧蜂属（Halictus sp．）和地蜂属（Andrena sp．）的2种昆虫是柴黄姜的主要传粉昆虫。访花昆虫对雄株的访花高峰时段为11：00至14：00,对雌花没有明显的访花高峰时段。在自然条件下,柴黄姜的座果率为18．86％,结实率为7．12％,种子萌发率为64．76％。在武当山,柴黄姜种群扩展仍以有性繁殖为主,无性繁殖为重要补充。     

番荔枝科两种植物花器官形态发生

用扫描电镜观察了囊瓣木（Saccopetalum prolificum）和刺果番荔枝（Annona muricata）花器官的形态发生过程。刺果番荔枝和囊瓣木花被片均为3轮,其中刺果番荔枝内轮花被片数目为3枚、5枚或7枚。囊瓣木花原基最初为圆锥形,最外轮3枚花被片很快发生,之后中、内轮花被片原基连续发生,3轮花被片互生。此时花原基为六边形。花被片分化完成时,圆球形雄蕊原基沿六边形花原基的6个边螺旋向心发生,最终近轮状排列于花原基上。刺果番荔枝的雄蕊较多（约1000枚）,首先在中轮花被片所对的花原基边缘发生,之后大量雄蕊螺旋状发生。心皮分化的早期阶段,与雄蕊原基很相似,当心皮数目逐渐增多时,不能分辩出发生的顺序。成熟花中,心皮和雄蕊全都被毛覆盖,毛具有粘住传粉滴的作用。     

Floral Variation in Tepals,Sterile and Fertile Stamens of Kingdonia uniflora (Ranunculaceae) with Reference to Pollinators and Pollination

To understand the influence of different floral organs of Kingdonia uniflora to the pollinators and pollination, we divided the flowers into four groups, A, with the sterile stamens removed, B, with the tepals removed, C, with the fertile　stamens removed and D, without the remove of any organ. The results showed that the insects visiting frequency were 0.4 time h in group A, 0 time h in group B, 0. 9 time h in group C and 2. 2 time h in group D. The percentage of pollinated flowers was 29.09% in group A, 40. 38% in group B, 70.91% in group C and 91. 67% in group D. The percentage of pollinated carpels was 17.77% in group A, 20.94% in group B, 40.58% in group C and 75.27% in group D respectively. The relationships between the insect visiting frequency and the nectarial secretion of the sterile stamens were observed in the field. To compare with group D, the decrease of the rate of pollinated flower and the carpel was group A > group B>group C. To combine the observation of the relationships between the nectarial secretion of the sterile stamens and the insect visiting frequency, we considered that the sterile stamens with nectar as the reward of the visiting insects play the most important rule for the pollinators and pollination. However, the tepals that could afford the platform of staining and moving of the insects on the flowers are also important for the pollinators. The fertile stamens that could form the chromatic to attract some of the pollinators and supply the pollen grains as the food of some pollinators play less important rule .     

舌瓣鼠尾草退化杠杆雄蕊的相关花部特征及传粉机制

杠杆状雄蕊是鼠尾草属(Salvia)物种形成的关键性状, 背部杠杆传粉模式作为该属植物与传粉者精确互作的经典案例被广泛深入研究, 但是在该属物种中还存在许多非典型的杠杆结构和传粉模式。雄蕊结构及其与传粉者互作的多样性, 使得鼠尾草属成为研究植物传粉模式转变的模式材料, 舌瓣鼠尾草(S. liguliloba)即是一种具非典型的退化杠杆状雄蕊结构和传粉特征的代表性物种。该文着重对舌瓣鼠尾草的花器官结构和传粉特征进行研究, 并与具有短药隔杠杆的毛地黄鼠尾草 (S. digitaloides)做比较分析, 以期揭示退化杠杆可能的进化选择压力及其生态学意义。结果表明, 舌瓣鼠尾草具有较短的花冠、更窄的冠筒和较短的雄、雌蕊(p < 0.05)。退化萎缩的雄蕊下臂, 冠筒内的狭小空间限制了唯一的有效传粉昆虫——三条熊蜂(Bombus trifasciatus)推动雄蕊做杠杆状运动, 而是靠近花药直接利用头部完成授粉。相比经典的杠杆状雄蕊结构及其传粉过程, 小型花冠和退化杠杆雄蕊是对专一性和活跃度较高传粉昆虫的适应, 可能具有完全不同的进化途径和繁殖策略。     

Fruit production in cranberry (Ericaceae: Vaccinium macrocarpon): a bet-hedging strategy to optimize reproductive effort

In the cultivated cranberry (Vaccinium macrocarpon), reproductive stems produce 1-3 fruit even though they usually have 5-7 flowers in the spring. We undertook experiments to test the hypothesis that this was an adaptive life history strategy associated with reproductive effort rather than simply the result of insufficient pollination. We compared fruit production on naturally pollinated plants with those that were either manually pollinated or that were caged to exclude insects. Clearly, insects are necessary for the effective pollination of cranberry plants, but hand pollination of all flowers did not result in an increase in fruit number. Most of the upper flowers, which had significantly fewer ovules than did the lower flowers, aborted naturally soon after pollination. However, when the lower flower buds were removed, the upper flowers produced fruit. This suggests that the upper flowers may serve as a backup if the earlier blooming lower ones are lost early in the season. Furthermore, the late-blooming flowers may still contribute to the plant's reproductive success as visiting pollinators remove the pollen, which could serve to sire fruit on other plants. These results are discussed in the context of their possible evolutionary and proximate causes.     

Mixed bumblebee and blowfly pollination of Cypripedium flavum (Orchidaceae) in Sichuan,China

Most Cypripedium species are specialized orchids pollinated by, in a broad sense, bees or flies. Here we present the first evidence that a slipper orchid, Cypripedium flavum, is pollinated by both bees and flies, i.e., bumblebees and blowflies. Artificial pollination experiments demonstrated that the flowers of C. flavum are self-compatible, but need pollen vectors for successful reproduction. Field observations detected 25 insects visiting the flowers, and 14 of these insects entered into the labellum of the flowers, but only female bumblebees, Bombus hypnorum, B. remotus, and the blowfly Calliphora vomitoria exited of the labellum with pollen smears of C. flavum. The floral functional morphology of C. flavum appears to be more suited to bumblebees than to blowflies. The bumblebees are more efficient pollinators of the orchid, but blowflies are more frequent visitors, so they pollinated more flowers despite being less efficient.     

Distribution of calcium oxalate crystals in floral organs of Araceae in relation to pollination strategy

? Premise of the study: Many flowers are pollinated by potentially hungry insects, yet flowers also contain gametes and embryos which must be protected from predation. Microscopic calcium oxalate crystals in plant tissues have been proposed to protect against herbivory. Aroids, which have an unusual diversity of such crystals, also exhibit diverse pollination strategies. Many species have pollinators that do not feed while visiting the flowers, while other species, especially those pollinated by beetles, offer sterile staminodia as food rewards. We examined flowers of 21 aroid species with various pollination strategies to test the hypothesis that crystals protect vital gametes and embryos while allowing consumption of food bribes. ? Methods: Aroid inflorescences collected from the field or from greenhouse material were sectioned, cleared, and examined by bright field and polarization microscopy. ? Key results: All species examined, regardless of pollination strategy, arrayed crystals around unshed pollen and ovules. Less vital tissues, such as odoriferous appendages, had few crystals. Staminodia offered as food to beetle pollinators, however, differed greatly between species in their crystal contents. Some had minimal crystals; some had crystals in patterns suggesting they limit beetle feeding; still others had abundant crystals in no obvious pattern. ? Conclusions: The results are consistent with crystals protecting against insect predation of gametes and embryos. However, the role of crystals in food-bribe staminodia is unclear. They may limit and direct feeding by beetles in some species, while in others they might have no protective role.     

High ratio of illegitimate visitation by small bees severely weakens the potential function of heteranthery

Aims In heterantherous plants, 'division of labor' among structurally different stamens, i.e. pollinating and feeding functions, has been thought to reduce the evolutionary conflict of relying on pollen both as the carrier of male gametes and as the food for pollinators. The key to successful division of labor among different sets of stamens is the size match between stamens and legitimate pollinators, which results in the precise deposition of pollen onto specific locations on pollinator's body and facilitates cross pollination. However, the potential impact of small illegitimate insects that are ubiquitous during the pollination process on the plant reproduction in heterantherous species has been largely neglected in previous studies and never been demonstrated experimentally.Methods Here, we investigated the functions of three different types of stamens in Commelina communis. The pollinator visitation, pollen removal and deposition were compared among flowers with different types of anthers emasculated at two natural populations. Moreover, the mating systems of C. communis in wild populations were estimated using microsatellite markers.Important findings Our data showed that the main floral visitors for C. communis at the two studied populations were small illegitimate bees rather than legitimate pollinators, accounting for 77.5 and 92.2% of total flower visits, respectively. Flower manipulations in C. communis demonstrated that the two types of brightly yellow stamens separately functioned as 'deceptive attraction' and 'feeding' functions. Although the brown inconspicuous stamens of C. communis with the largest amount of fertile pollen had the potential function in offering pollen for cross pollination, the high ratio of illegitimate visitation by small bees significantly affected the dispersal and deposition of pollen from the pollinating anthers, and subsequently decreased the levels of outcrossing (t m = 0.23–0.32) in wild populations. Our work further confirmed that the size match between pollinators and the floral morphology is the prerequisite to successfully fulfill the functional differentiation among different sets of stamens in heterantherous plants. Local high ratio of illegitimate visitation by size unmatched insects could significantly weaken the potential functions of heteranthery, affecting the dispersal and deposition of functional pollen in heterantherous plants and further the whole mating systems.     

Mechanisms that limit pollinator range in Ericaceae

Studies were conducted in 2001-2003 at Valdai National Park (Novgorod region) and at the Zvenigorod biological station of Moscow State University. The morphology of flowers, flowering dynamics and composition of insect visiting flowers of Ericaceae species: Andromeda polifolia, Chamaedaphne calyculata, Ledum palustre, Oxycoccus palustris, Vaccinium myrtillus, V. uliginosum, and V. vitis-idaea L. were studied. Some species of insects visiting flowers were excluded from the list of pollinators on the basis of observation on their behavior. L. palustre was visited mainly by flies where as other investigated species were visited mainly by bumblebees. In some cases bumblebees were the only visitors of the investigated plants. Mechanisms that protect flowers from flies and short-tongued solitary bees visits and ensure a best pollination by bumblebees are various among different species of Ericaceae. Efficiency of nectary protection also differs among different plant species and is defined by particularities of their habitats and flowering phenology. As far as all species of this family during the flowering are dominants in typical habitats, a competition for the pollination with species of other families in most cases is megligible. Flowering periods of V. vitis-idaea and V. myrtillus in forest ecosystems overlapped weakly. Moreover, V. myrtillus is pollinated mainly by bumblebee queens where as pollinators of V. vitis-idaea are bumblebee workers, solitary bees and horse flies. The other investigated plant species inhabit only oligotrophic peat bogs. Thery are pollinated by bumblebees but periods of flowering are not overlapped and consequently follow one after another. L. palustre and V. uliginosum flower simultaneosly but they are pollinated by different pollinators.     

两性花的雄蕊运动:多样性和适应意义

雄蕊运动指雄蕊在自身能量支持下发生的主动运动,不包括雄蕊在访花者触碰下造成的被动位移。该文总结了雄蕊的应激运动、快速猛烈弹射、缓慢运动以及级联运动等4种主要类型,分析了这些运动类型的系统分布及繁殖适应意义等方面的研究进展。雄蕊的应激运动由访花者或其他外力诱发,可能起到促进散粉和实现繁殖保障的作用;雄蕊快速猛烈的弹射运动可将花粉猛然撒向空中或访花者身上,促进了花粉的风媒或虫媒散布;缓慢运动的雄蕊可能通过在不同花期改变雄蕊的空间位置和雌雄异位程度来调节繁殖策略,或主动将雄蕊花药移至特定部位(如柱头表面)实现自交;雄蕊逐一、依次发生的级联运动较为复杂,主要分布在刺莲花科、梅花草科、旱金莲科和芸香科中,目前还缺乏实验研究;但根据"花粉呈现理论"以及其他类型的雄蕊运动研究结果,雄蕊的级联运动可以将花粉分批呈现给不同的传粉者,通过不同传粉者的分别传粉来提高花粉的输出;而且可避免已散粉雄蕊对即将散粉雄蕊的干扰,同时可能也降低了雌雄功能干扰和(或)花内自交。在芸香(Ruta graveolens)中,级联运动之后的雄蕊还会在花末期再同时向花中央运动;这种多向、多次运动方式是目前发现的最复杂的雄蕊运动类型。雄蕊运动领域值得今后开展进一步实验研究的方向主要有:1)雄蕊运动尤其是级联运动对雌雄功能干扰(性别间干扰)、雄蕊与雄蕊的"性别内干扰"等植物繁殖格局的影响;2)雄蕊运动与雌雄异熟、雌雄异位等花部特征的相互作用;3)雄蕊运动复杂类型的生理与发育机制。