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Siphonogamy, the delivery of nonmotile sperm to the egg via a pollen tube, was a key innovation that allowed flowering plants (angiosperms) to carry out sexual reproduction on land without the need for water. This process begins with a pollen grain (male gametophyte) alighting on and adhering to the stigma of a flower. If conditions are right, the pollen grain germinates to produce a pollen tube. The pollen tube invades the stigma and grows through the style towards the ovary, where it enters an ovule, penetrates the embryo sac (female gametophyte) and releases two sperm cells, one of which fertilizes the egg, while the other fuses with the two polar nuclei of the central cell to form the triploid endosperm. The events before fertilization (pollen-pistil interactions) comprise a series of complex cellular interactions involving a continuous exchange of signals between the haploid pollen and the diploid maternal tissue of the pistil (sporophyte). In recent years, significant progress has been made in elucidating the molecular identity of these signals and the cellular interactions that they regulate. Here we review our current understanding of the cellular and molecular interactions that mediate the earliest of these interactions between the pollen and the pistil that occur on or within the stigma - the 'pollen-stigma interaction'.

To minimize interspecific pollination, it has been suggested that pollen is placed on different parts of a pollinator's body corresponding to the conspecific location of pollen pickup by the stigma. Although Pedicularis is regarded as a classic example of pollinator‐mediated floral isolation, such reciprocal pollen placement has not been demonstrated experimentally. This leads us to question previous observations of pollen release in Pedicularis species. Here, we show that pollen grains are released from the tip, rather than the basal opening, of the galea (the hoodlike upper lip of the corolla) in eight nectarless Pedicularis species, mimicking pollen release from poricidal anthers. We used safranin‐stained pollen within anthers to track pollen placement in three Pedicularis species, and showed that pollen was deposited on numerous parts of the bumblebee's body. However, fluorescent powder placed on the stigmas to detect the contact location on the bumblebee's body was deposited mainly on the major position of pollen placement in each of the three species. Such segregation of pollen placement and pickup between species sharing the same pollinator probably helps to reduce reproductive interference, but the positions of pollen placement and stigma contact on the bumblebee's body were not as precise as previously thought.

Community studies have shown that plant species are often pollinated by multiple pollinators; however, networks of heterospecific pollen transfer (HPT) in natural communities remain largely unexplored. We analyzed pollen deposition on stigmas of 57 flowering species to build a picture of plant-plant interactions via HPT in a biodiverse alpine meadow in southwest China. Plant species were categorized as pollen donors or recipients by their link numbers and link qualities. We identified 3609 heterospecific pollen grains, representing 410 links among 69 pollen species. Each plant species received on average 7.2 pollen species and donated its pollen to 5.5 species; only a few species donated or received large amounts of pollen or pollen from a large number of species. Compared to specialized plants, generalized plants tended to receive more heterospecific pollen but exported no more pollen to other species. Plant position in the network was related to both floral traits (stigma position) and pollinator generalization level. When different species share the same pollinator, bidirectional HPT may occur, but this was rarely observed in the species-rich community, indicating that interspecific pollen interference was largely unidirectional. Our study highlights the importance of understanding how sympatric flowering plants reduce deleterious effects of HPT, for example via stigma position. This study is the first to present a pollen transfer network for an entire community and to unravel its properties using directed network analysis.

Structural organization of organs in multicellular organisms occurs through intricate patterning mechanisms that often involve complex interactions between transcription factors in regulatory networks. For example, INDEHISCENT (IND), a basic helix-loop-helix (bHLH) transcription factor, specifies formation of the narrow stripes of valve margin tissue, where Arabidopsis thaliana fruits open on maturity. Another bHLH transcription factor, SPATULA (SPT), is required for reproductive tissue development from carpel margins in the Arabidopsis gynoecium before fertilization. Previous studies have therefore assigned the function of SPT to early gynoecium stages and IND to later fruit stages of reproductive development. Here we report that these two transcription factors interact genetically and via protein-protein contact to mediate both gynoecium development and fruit opening. We show that IND directly and positively regulates the expression of SPT, and that spt mutants have partial defects in valve margin formation. Careful analysis of ind mutant gynoecia revealed slight defects in apical tissue formation, and combining mutations in IND and SPT dramatically enhanced both single-mutant phenotypes. Our data show that SPT and IND at least partially mediate their joint functions in gynoecium and fruit development by controlling auxin distribution and suggest that this occurs through cooperative binding to regulatory sequences in downstream target genes.

In the Brassicaceae, compatible pollen-pistil interactions result in pollen adhesion to the stigma, while pollen grains from unrelated plant species are largely ignored. There can also be an additional layer of recognition to prevent self-fertilization, the self-incompatibility response, whereby self pollen grains are distinguished from nonself pollen grains and rejected. This pathway is activated in the stigma and involves the ARM repeat-containing 1 (ARC1) protein, an E3 ubiquitin ligase. In a screen for ARC1-interacting proteins, we have identified Brassica napus Exo70A1, a putative component of the exocyst complex that is known to regulate polarized secretion. We show through transgenic studies that loss of Exo70A1 in Brassica and Arabidopsis thaliana stigmas leads to the rejection of compatible pollen at the same stage as the self-incompatibility response. A red fluorescent protein:Exo70A1 fusion rescues this stigmatic defect in Arabidopsis and is found to be mobilized to the plasma membrane concomitant with flowers opening. By contrast, increased expression of Exo70A1 in self-incompatible Brassica partially overcomes the self pollen rejection response. Thus, our data show that the Exo70A1 protein functions at the intersection of two cellular pathways, where it is required in the stigma for the acceptance of compatible pollen in both Brassica and Arabidopsis and is negatively regulated by Brassica self-incompatibility.

1. The honeybee Apis mellifera is currently in decline worldwide because of the combined impacts of Colony Collapse Disorder and the Varroa destructor mite. In order to gain a balanced perspective of the importance of both wild and managed pollination services, it is essential to compare these services directly, a priori, within a cropping landscape. This process will determine the capacity of other flower visitors to act as honeybee replacements. 2. In a highly modified New Zealand agricultural landscape, we compared the pollination services provided by managed honeybees to unmanaged pollinator taxa (including flies) within a Brassica rapa var. chinensis mass flowering crop. 3. We evaluate overall pollinator effectiveness by separating the pollination service into two components: efficiency (i.e. per visit pollen deposition) and visit rate (i.e. pollinator abundance per available flower and the number of flower visits per minute). 4. We observed 31 species attending flowers of B. rapa. In addition to A. mellifera, seven insect species visited flowers frequently. These were three other bees (Lasioglossum sordidum, Bombus terrestris and Leioproctus sp.) and four flies (Dilophus nigrostigma, Melanostoma fasciatum, Melangyna novae-zelandiae and Eristalis tenax). 5. Two bee species, Bombus terrestris and Leioproctus sp. and one fly, Eristalis tenax were as efficient as the honeybee and as effective (in terms of rate of flower visitation). A higher honeybee abundance, however, resulted in it being the more effective pollinator overall. 6. Synthesis and applications. Alternative land management practices that increase the population sizes of unmanaged pollinator taxa to levels resulting in visitation frequencies as high as A. mellifera, have the potential to replace services provided by the honeybee. This will require a thorough investigation of each taxon's intrinsic biology and a change in land management practices to ensure year round refuge, feeding, nesting and other resource requirements of pollinator taxa are met.

In natural communities, plants can receive pollen from multiple heterospecifics as well as conspecifics. However, studies on the effects of interspecific pollen transfer have focused on interactions between species pairs. The potential exists for diverse interactions among heterospecific pollen (HP) grains on the stigma, and for these to affect plant reproduction, alone or in combination with conspecific pollen (CP) loss, but these interactions have not yet been explored. We used hand-pollinations to simulate increasing community diversity and CP loss on Mimulus guttatus stigmas. We used pollen mixes of one to three heterospecific donors to determine how species composition and CP load size affect seed production and to characterize the mechanisms underlying fertilization failure. Heterospecific pollen deposition reduced M. guttatus seed production and while the effect increased with the number of heterospecific donors, the strength depended on species composition and was independent of conspecific load size. Different types of interactions (additive and synergistic) are hypothesized to underlie the diverse effects on M. guttatus reproductive success. Our results suggest that an increase in the diversity of heterospecific donors will not always lead to a greater decrease in fitness because multispecies effects depend on the interacting species.

Background and AimsInterest in pollinator-mediated evolutionary divergence of flower phenotype and speciation in plants has been at the core of plant evolutionary studies since Darwin. Specialized pollination is predicted to lead to reproductive isolation and promote speciation among sympatric species by promoting partitioning of (1) the species of pollinators used, (2) when pollinators are used, or (3) the sites of pollen placement. Here this last mechanism is investigated by observing the pollination accuracy of sympatric Pedicularis species (Orobanchacae).MethodsPollinator behaviour was observed on three species of Pedicularis (P. densispica, P. tricolor and P. dichotoma) in the Hengduan Mountains, south-west China. Using fluorescent powder and dyed pollen, the accuracy was assessed of stigma contact with, and pollen deposition on, pollinating bumble-bees, respectively.Key ResultsAll three species of Pedicularis were pollinated by bumble-bees. It was found that the adaptive accuracy of female function was much higher than that of male function in all three flower species. Although peak pollen deposition corresponded to the optimal location on the pollinator (i.e. the site of stigma contact) for each species, substantial amounts of pollen were scattered over much of the bees' bodies.ConclusionsThe Pedicularis species studied in the eastern Himalayan region did not conform with Grant's ‘Pedicularis Model’ of mechanical reproductive isolation. The specialized flowers of this diverse group of plants seem unlikely to have increased the potential for reproductive isolation or influenced rates of speciation. It is suggested instead that the extreme species richness of the Pedicularis clade was generated in other ways and that specialized flowers and substantial pollination accuracy evolved as a response to selection generated by the diversity of co-occurring congeners.

BACKGROUND AND AIMS: From Darwin's time onward, biologists have thought about adaptation as evolution toward optimal trait values, but they have not usually assessed the relative importance of the distinct causes of deviations from optima. This problem is investigated here by measuring adaptive inaccuracy (phenotypic deviation from the optimum), using flower pollination as an adaptive system. METHODS: Adaptive accuracy is shown to have at least three distinct components, two of which are optimality (deviation of the mean from the optimum) and precision (trait variance). We then describe adaptive accuracy of both individuals and populations. Individual inaccuracy comprises the deviation of the genotypic target (the mean phenotype of a genotype grown in a range of environments) from the optimum and the phenotypic variation around that genotypic target (phenotypic imprecision). Population inaccuracy has three basic components: deviation of the population mean from the optimum, variance in the genotypic targets and phenotypic imprecision. In addition, a fourth component is proposed, namely within-population variation in the optimum. These components are directly estimable, have additive relationships, and allow exploration of the causes of adaptive inaccuracy of both individuals and populations. Adaptive accuracy of a sample of flowers is estimated, relating floral phenotypes controlling pollen deposition on pollinators to adaptive optima defined as the site most likely to get pollen onto stigmas (male inaccuracy). Female inaccuracy is defined as the deviation of the position of stigma contact from the expected location of pollen on pollinators. KEY RESULTS: A surprising amount of variation in estimated accuracy within and among similar species is found. Some of this variation is generated by developmental changes in positions of stigmas or anthers during anthesis (the floral receptive period), which can cause dramatic change in accuracy estimates. There seem to be trends for higher precision and accuracy in flowers with higher levels of integration and dichogamy (temporal separation of sexual functions), and in those that have pollinators that are immobile (or immobilized) during pollen transfer. Large deviations from putative adaptive optima were observed, and these may be related to the effects of conflicting selective pressures on flowers, such as selection against self-pollination promoting herkogamy (spatial separation of pollen and stigmas). CONCLUSIONS: Adaptive accuracy is a useful concept for understanding the adaptive significance of phenotypic means and variances of floral morphology within and among populations and species. Estimating and comparing the various components of adaptive accuracy can be particularly helpful for identifying the causes of inaccuracy, such as conflicting selective pressures, low environmental canalization and developmental instability.