Pollination system of Holcoglossum rupestre (Orchidaceae): A special and unstable system

The pollination system of Holcoglossum rupestre was studied in northwestern Yunnan. The species is self-compatible but pollinator-dependent and offers nectar as a reward to visitors. A species of beetle, Hybovalgus bioculatus Kolbe, is identified as the pollinator among all visitors observed. The beetles carried the pollinia in two different positions, i.e. on the buttock and the foreleg, and pollinated the flowers by two different mechanisms. The analysis suggests that Holcoglossum rupestre and its pollinator may be in an unstable pollination relationship. The natural fruit set is low, which is attributed to the low visiting frequency of pollinator and the effective mechanism to prevent self-pollination.     

Reproductive biology of South American Bothriochloa (Poaceae: Andropogoneae)

The reproductive biology of 13 South American taxa of Bothriochloa was studied. All surveyed species reproduce sexually and are self-compatible. Several elements also point to self-pollination as their main breeding system: (i) bud pollination that occurs in most species enhancing cleistogamy, (ii) presence of pits in the glumes that impede the opening of flowers and (iii) reduction in the number of stamens, a trend associated with outbreeding.     

蛛网萼开花物候、花部特征及繁育系统研究

为探讨蛛网萼(Platycrater arguta)传粉生物学特性,该研究以江西上饶广丰铜钹山国家级自然保护区、江西阳际峰国家级自然保护区蛛网萼为材料,采用野外观察和人工授粉等方法,对蛛网萼开花物候、花部形态及繁育系统进行了研究。结果表明:(1)蛛网萼为伞形花序,种群花期一般在6月中下旬至8月中旬,单花花期为2~3 d,花序花期持续10 d。两个保护区由于年均温度与光照的不同,花期进程有显著不同。(2)蛛网萼可孕花与不孕花从绿色变为白色时对访花昆虫有吸引作用,不孕花能够增加昆虫的访花频率。(3)蛛网萼花粉胚珠比(P/O)为2 520~3 100,杂交指数即是3或4。(4)繁育体系属于以异交为主,部分自交亲和,需要传粉者;熊蜂(Bombus sp.)、四条蜂(Apis florea)和油茶地蜂(Andrena cammelia)是蛛网萼的有效传粉昆虫。(5)套袋和人工授粉结果显示,蛛网萼为兼性自交和异交授粉,异株异花授粉能显著提高坐果率及结籽率,这主要与生境中有效传粉昆虫种类、数量少,花粉传递效率低,种群间基因流交流降低以及柱头自花及同株异花传粉等有关,从而表明在蛛网萼现有生境中植株的早期生殖成功受到影响。蛛网萼种群分布范围扩张以有性繁殖为主,无性繁殖作为重要补充及繁殖保障。     

紫花含笑传粉生物学初探

紫花含笑(Michelia crassipes)花色艳丽、花香浓郁,为解释其在自然条件下结实率低的原因和判断最佳杂交时期,该研究从传粉生物学入手,通过观察紫花含笑开花动态,运用异交指数(OCI)、花粉胚珠比(P/O)、花粉和柱头活性、人工授粉试验等方法对其繁育系统进行了检测,并记录访花昆虫的种类和行为。结果表明:(1)紫花含笑花期在4—5月,持续50 d左右,单花花期4~5 d。(2)雌雄同花,雌蕊先成熟,雄蕊空间位置高于雌蕊。(3)OCI等于4,P/O为2671±123。(4)套袋实验表明紫花含笑能进行少量的自花传粉,人工异花授粉结实率和种子质量显著高于人工自花授粉和自然条件下,且不存在无融合生殖。(5)开花后36 h雌蕊群弯曲分泌粘液时,与含笑(M.figo)杂交结实率最高达85.7%,为最佳授粉时期。(6)有效访花者主要有东方蜜蜂、黑纹食蚜蝇和独角仙,访花频率易受天气影响。综上说明,紫花含笑繁育系统以异交为主,部分自交亲和需要传粉者,存在一定近交衰退,自然条件下结实率低,主要受传粉昆虫和雌雄异熟限制,最佳授粉期的确定能够提高其杂交成功率。该研究结果为紫花含笑资源保护和杂交育种奠定了基础。     

The occurrence of estrogens in relation to reproductive processes in flowering plants

Summary Estrogens, a group of steroidal sex hormones, were determined in pollen, styles or whole pistils using a highly sensitive and specific radioimmunoassay (RIA) and found to be present at levels between 10^–11 and 10^–10 g/g FW. In the style of Liliun davidii Duch., the levels of total estrogen decreased after self-pollination of the open flower. After bud pollination, which partly overcame incompatibility, the levels of both estradiol and total estrogen increased. The levels of estradiol and total estrogen in the pistils of Brassica pekinensis Rupr. increased after compatible pollination during the first 3 days following pollination and decreased thereafter when the data were expressed on a FW basis. These results suggest that estrogens are to some extent associated with the reproductive processes.     

A supplementary contribution of ants in the pollination of an orchid, Epipactis thunbergii, usually pollinated by hover flies

It has been controversial how extensively ants contribute to pollination, and we evaluated the contribution of the Japanese carpenter ant, Camponotus japonicus, to the pollination of an orchid, Epipactis thunbergii. Two-year field studies revealed that (1) the ant workers foraged even in cool/cloudy conditions and accordingly visited orchid flowers more frequently (about 40% of all the visitors) than hover flies, the principle pollinators (10–20%), and that (2) the flower-visiting ants occasionally removed pollinia from the anther and then delivered pollen onto the stigmatic surface of other flowers, although self-pollination might frequently occur in the consecutive visits of flowers within an inflorescence. An artificial pollination experiment with pollinia which had been transferred to the ant integument showed that (3) the treated flowers produced as many fruits and seeds as control flowers. We concluded that C. japonicus workers could actually pollinate E. thunbergii flowers and their relative importance as pollinators appeared to be largely dependent on the abundance of flower-visiting workers or weather conditions during the flowering period, which mainly determined the availability of hover flies.     

Quantitatively determined self-incompatibility

Summary It has been claimed that Borage (Borago officInalis L.) has a multifactorial self-incompatibility system. Such systems may have a high level of ineffective pollination, and we show that this is the case in borage. The ranking of seed set from highest to lowest is as follows: bee-pollination; natural pollination in the absence of bees; artificial cross-pollination between unrelated plants; artificial cross-pollination between related plants; artificial self-pollination. In diallel crosses, significant parental effects were detected but no consistent patterns of seed set, which suggest a simple self-incompatibility system, were detected. The level of outcrossing with natural pollination was very variable but greater than 50%. Thus, there appears to be no straightforward self-incompatibility system in borage.     

Rates of cross-pollination inLycopersicon pimpinellifolium: Impact of genetic variation in floral characters

Cross-pollination was tested between different genotypes ofLycopersicon pimpinellifolium in an experimental plot situated in the southerly range of the species. Rates of cross-pollination were found to be highly correlated with flower size and were related to a lesser extent with degree of stigma exsertion. Comparisons of fertility suggest that exsertion of 1.5 mm or more from the anther tube orifice interferes with automatic self-pollination, thereby increasing rates of cross-pollination. A single species ofExomalopsis (Anthophoridae), observed to visit all tested genotypes ofL. pimpinellifolium, was responsible for most of the pollination. Other species of solitary bees and other bee families participated in cross-pollination.     

Differentiated protein synthesis with polysomes from Petunia ovaries before fertilization

Polysomes from cross-, self- and unpollinated ovaries have been injected in Xenopus laevis eggcells. The synthesised proteins have been identified by means of gel-electrophoresis. It appeared that few hours after pollination the protein pattern of the pollinated ovary has changed in comparison with that of unpollinated ones. However, in this early stage, there is only little difference found in the protein pattern of selfed and crossed ovaries. From 12 h after pollination and onwards, the difference between pollinated and unpollinated ovaries becomes very obvious, as does also the difference between crossed and selfed ovaries. At 24 h after pollination many different proteins exist in crossed and selfed ovaries. From 32 h to 48 h after pollination the crossed ovaries differentiate further on, whereas the ovaries after self-pollination show more and more proteins which were also found in the unpollinated ovaries of that age. Possible explanations and hypothesis for these phenomena are discussed as well as the probable occurrence of signal(s) arriving from the stigma or the style at the ovary.     

Pollination mechanism in southern Brazilian orchids which are exclusively or mainly pollinated by halictid bees

The pollination process is documented in three sympatric orchids found in Curitiba, Paraná State, southern Brazil. InCampylocentrum aromaticum (Epidendroideae: Angraeciinae) andPrescottia densiflora (Spiranthoideae: Prescotiinae) halictid bees act as main pollinators with minor contributions from other insects, such as Syrphid flies. Because of the pollinarium and column structure inCyclopogon diversifolius. (Spiranthoideae, Spiranthinae), halictid bees are the only insects capable of performing pollination. In the three studied species pollinarium removal and deposition are achieved by interaction between the column and bee's mouth parts. Morphological and ecological flower features in agreement with this pollination mode are discussed and pollination by halictid bees is suggested to occur in a number of South American orchids.     

Floral biology of Nuphar pumila (Timm) DC. (Nymphaeaceae) in China

Anthesis in individual flowers of Nuphar pumila (Timm) DC. occurs for four consecutive days. First-day flowers are protogynous and functionally female. Flowers can open completely on the first day of anthesis. This contrasts with all previous reports, which state that first-day flowers of Nuphar are characterized by partial expansion of the calyx, leaving a distal small triangular opening just above the stigmatic disc. Flowers close completely on the first night of anthesis and remain partially open on the subsequent three nights. During the entire anthesis period the stigmas emit a sweet, fruity odor and the petal nectaries produce visible nectar drops. The stigma of N. pumila is secretory and unicellular-papillate. Pollen grains are monosulcate with long spines. Our observations on the mating system of N. pumila indicate that neither asexual seed production nor spontaneous self-pollination occurs. Cross-pollination of second-, third- and fourth-day flowers produced few seeds. Flowers of N. pumila were mainly pollinated by sweat bees, with flies playing a minor pollination role. No beetle visits were observed. Our insect-pollination observations substantiate the view that the relative contribution of flies, bees, and beetles to pollination in a single Nuphar population depends on two factors: the relative abundance of the insects, and presence of alternative food sources.     

授粉方式对铁皮石斛种质座果及种子萌发的影响

为进行铁皮石斛(Dendrobium officinale)的良种选育和种苗生产,对46个铁皮石斛种质进行人工授粉试验,研究了不同授粉方式的授粉成功率、座果率和种子的萌发能力。结果表明,4种授粉方式的授粉成功率明显不同,不同种质间的授粉成功率可达100%,且座果率普遍较高。只有31.3%的种质可以成功自花授粉,且座果率相差较大。同株异花授粉和种质内异株授粉的成功率和座果率均介于上述二者之间。授粉后所得种子的萌发率和萌发速度在4种授粉方式间也呈现与授粉成功率和座果率一致的趋势。这表明在铁皮石斛种质中存在自交不亲和现象,因此以不同产地来源的铁皮石斛种质为亲本进行人工授粉,可大大提高授粉的成功率和座果率,以及后代种子的萌发能力。     

Pollination biology and breeding systems in the terrestrial orchid Bletilla striata

The levels of fruit set in different habitats (three inland populations vs. three coastal populations) and the breeding system in the terrestrial orchid Bletilla striata were examined in two groups of populations in Haenam-gun, Jollanam-do Province in southern Korea. In total inland populations (mean fruit set=13.3%) showed about eight-fold higher fruit set per plant than the coastal populations (1.6%), probably due to mild climatic conditions in the inland populations during the flowering period. Honey bee workers Apis mellifera were observed in the inland populations, but no visitors were found in the coastal populations. However, a great increase in fruit set was achieved by hand-pollination: the percentage of fruit set through artificial self-pollination and geitonogamous pollination were close to 90%, indicating that B. striata is self-compatible. Agamospermy and spontaneous autogamy (automatic self-pollination) were not detected in this study. These indicate that pollinia vectors are essential for achieving fruit set in natural populations of B. striata. Except in one inland population examined in 2001, there was no significant correlation between the patch size per population and the average percentage of fruit set, which might reflect deception-pollination of B. striata and/or paucity of pollinators. Overall levels of fruit sets within and among patches in each population were homogeneous during a two-year period.     

The interspecific hybrid Petunia parodii × P. inflata and its relevance to somatic hybridization in the genus Petunia

Summary Attempts at the reciprocal cross between Petunia parodii and P. inflata using standard emasculation and pollination techniques failed. Limited pollen tube growth down the style in reciprocal crosses led to reproductive isolation between the self-compatible P. parodii and self-incompatible P. inflata. The interspecific hybrid was successfully produced by bud-pollination of P. parodii with P. inflata as the male parent in 22 percent of attempts, but not in the opposite direction. In vitro pollination of P. parodii ovaries with P. inflata pollen also produced hybrids. The small size of the ovary made it technically impossible to use P. inflata as the female parent for in vitro pollination. The interspecific hybrids were intermediate, as compared to the two parents, for six of the seven plant and flower characters measured. Furthermore, the hybrids had high pollen fertility, set abundant seed upon self-pollination, and readily inter-crossed with the parental species. The results are consistent with a high degree of chromosomal homology in the parental species and with minor genetic divergency leading to reproductive isolation that is pre-zygotic in nature. Overcoming the barriers to cross-incompatibility by practical techniques resulted in fertile interspecific hybrids that segregated for parental characters. The potential value of employing the parental species in somatic hybridization experiments is discussed.Michigan Agricultural Experiment Station Journal Article No. 8404     

Reproductive biology of twoPrunus species (Rosaceae) in the Northwest Iberian Peninsula

We studied the reproductive biology ofPrunus spinosa andPrunus mahaleb (Prunoideae, Rosaceae) in the northwest Iberian Peninsula. The two species flowered at the same time (peaking on March 9 and 11, respectively in 1990) but differ significantly in their fruit maturation times. Nectar volume peaked in the early morning in both species, and was ten times greater inP. spinosa than inP. mahaleb. Neither species shows apomixis, nor does fruit-set occur if pollinators are excluded. In both species self-pollination resulted in fewer fruits than open pollination. The principal pollinators belong to theApidae family (79% and 63% of visits toP. spinosa andP. mahaleb, respectively). Results are compared with those for other rosaceous plants with fleshy fruits.     

Transfer of pollinaria on birds’ feet: a new pollination system in orchids

We report the discovery of a new mechanism of pollination in orchids: transfer of pollinaria on the feet of birds. Observations carried out in South Africa and Malawi showed that the orchids Disa chrysostachya Sw. and Disa satyriopsis Kraenzl. are pollinated by sunbirds. Pollinaria of these orchids become attached firmly to the birds toes when they perch on the tall narrow inflorescences which are packed tightly with numerous small orange flowers. Birds typically perch on the lower part of an inflorescence while reaching up to feed on nectar in flowers on the upper part, but occasionally reverse this position to probe the lower flowers. The nectar is contained within a short bulbous spur with a narrow entrance that permits entrance of a sunbirds slender tongue. Contrary to expectation, the pollination mechanism in D. chrysostachya is remarkably efficient with about 6.1% of pollen reaching stigmas on other plants and fruit set occurring in 95% of flowers at one site. Birds seldom move their feet once perched, thus minimizing the incidence of self-pollination, either within or between flowers on an inflorescence.     

Intrapopulation variation in the breeding system ofVillarsia lasiosperma (Menyanthaceae), a distylous species

The restricted perennial diploid herb,Villarsia lasiosperma, of southwestern Western Australia has distylous flowers with dimorphism in style and stamen length, stigma morphology, and pollen size. In order to assess the presence and nature of an incompatibility system in this species, a crossing program was carried out using 17 plants grown from seeds collected in two field populations. Pollen stainabilities of these plants mostly exceeded 95%. Mean seed-set of Longs following intermorph pollinations was c. 24 seeds per pollination and for Shorts was c. 16 seeds per pollination. Approximately three-quarters of the Longs and Shorts produced no seeds after self-pollination; the remainder produced very few seeds following such pollinations. No Shorts produced seeds after intramorph cross-pollinations. However, about half the Longs exhibited various levels of weakened intramorph incompatibility, with mean seed-sets following intramorph cross-pollinations up to half the mean seed-sets following intermorph pollinations. In the behavior of Longs, the incompatibility system ofV. lasiosperma differs from the idealized distylous breeding system, but resembles that of a purported close relative,V. exaltata of eastern and southeastern Australia.     

Limitations on reproductive success in endemic <Emphasis Type="Italic">Aquilegia viscosa</Emphasis> (Ranunculaceae) relative to its widespread congener <Emphasis Type="Italic">Aquilegia vulgaris</Emphasis>: the interplay of herbivory and pollination

Plant reproduction can be strongly affected by herbivory and different features of pollination ecology, such as pollinator visitation rates and capacity for self-pollination. The purpose of this study is to compare the relative impact of herbivory and pollination on maternal reproductive success in endemic Aquilegia viscosa and its widespread congener Aquilegia vulgaris. We conducted herbivore exclusion experiments in two populations of each species in 2 different years and showed that the maternal fertility of A. viscosa was significantly more limited by floral predation and pre-dispersal seed predation than its widespread congener. In the absence of herbivory, A. viscosa retained significantly lower maternal fertility than A. vulgaris. Experimental pollinations in an insect-free glasshouse showed that the two species have an equal seed/ovule ratio both in the absence of pollinators and in the presence of non-limiting outcross pollination. Pollinator visitation rates were significantly higher in populations of A. vulgaris than in populations of A. viscosa. In addition, path analyses showed that spur length, an important trait for pollinator attraction in Aquilegia, and, indirectly sepal and petal width, contribute positively to the seed/ovule ratio in A. vulgaris, but not in A. viscosa. These results indicate that maternal fertility of endemic A. viscosa is strongly reduced by flower and seed predation despite low rates of pollinator visitation, and that pollen or resource limitation in the wild may further reduce maternal fertility. Finally, floral trait variation appears to be decoupled from fertility variation in endemic A. viscosa, which possibly constrains the evolution of reproductive traits in this species.     

Temporal variation in pollinarium size after its removal in species of Bulbophyllum: A different mechanism preventing self-pollination in Orchidaceae

InBulbophyllum involutum andB. ipanemense (Orchidaceae), two closely related species, shortly after removal the pollinarium has a diameter of approximately twice that of the entrance of the stigmatic cavity, requiring a mean time of 105 to 135 minutes to shrink in width and allow pollination. Because the pollinators of these species remain for some minutes in the same flower after removing the pollinia this mechanism, previously unknown inOrchidaceae, may be very important in preventing self-pollination. This mechanism does not occur inB. weddellii, and the pollinator does not remain in the flower after removing the pollinia. The smaller diameter of the stigmatic cavity inB. involutum reduces by 50% the chances of interspecific pollination withB. weddellii, and interspecific crossing is strictly unidirectional. This is important in maintaining isolation between these sympatric species, which share the same pollinators and have synchronized flowering.     

Reproductive biology ofAsphodelus aestivus (Asphodelaceae)

InAsphodelus aestivus the large number of showy flowers opened per inflorescence, high nectar and pollen production, and absence of automatic self-pollination, so that insect visitation is necessary for pollination, indicate that cross-fertilization is favoured. LargeHymenoptera are the main pollinators. Hand self-pollinations result in some degree of fruit-set, but cross-pollination also yield low fruit- and seed-set. The perennial habit and polyploid status are in agreement with the breeding system of this species, with sexual reproduction being rather inefficient and active vegetative propagation.