A Note on the Evolution of the Stamens in the Laurales,with Emphasis on the Lauraceae

The number of pollen sacs per anther and their position have traditionally been among the most important systematic characters in the Lauraceae. Opinions differ, however, about the evolutionary direction in these characters. This paper is intended to draw attention to little known morphological features of the stamens, in particular to transitions between disporangiate and tetrasporangiate anthers. In addition, the phylogenetic derivation of the stamen appendages is discussed. It is hypothesized that branched androecial structures may be primitive within the Laurales, if not in the angiosperms as a whole.     

Light controls stamen elongation via cryptochromes,phytochromes and COP1 through HY5 and HYH

In Arabidopsis, stamen elongation, which ensures male fertility, is controlled by the auxin response factor ARF8, which regulates the expression of the auxin repressor IAA19. Here, we uncover a role for light in controlling stamen elongation. By an extensive genetic and molecular analysis we show that the repressor of light signaling COP1, through its targets HY5 and HYH, controls stamen elongation, and that HY5 – oppositely to ARF8 – directly represses the expression of IAA19 in stamens. In addition, we show that in closed flower buds, when light is shielded by sepals and petals, the blue light receptors CRY1/CRY2 repress stamen elongation. Coherently, at flower disclosure and in subsequent stages, stamen elongation is repressed by the red and far‐red light receptors PHYA/PHYB. In conclusion, different light qualities – sequentially perceived by specific photoreceptors – and the downstream COP1–HY5/HYH module finely tune auxin‐induced stamen elongation and thus male fertility.     

ATP binding cassette transporters ABCG1 and ABCG16 affect reproductive development via auxin signalling in Arabidopsis

Angiosperm reproductive development is a complex event that includes floral organ development, male and female gametophyte formation and interaction between the male and female reproductive organs for successful fertilization. Previous studies have revealed the redundant function of ATP binding cassette subfamily G (ABCG) transporters ABCG1 and ABCG16 in pollen development, but whether they are involved in other reproductive processes is unknown. Here we show that ABCG1 and ABCG16 were not only expressed in anthers and stamen filaments but also enriched in pistil tissues, including the stigma, style, transmitting tract and ovule. We further demonstrated that pistil‐expressed ABCG1 and ABCG16 promoted rapid pollen tube growth through their effects on auxin distribution and auxin flow in the pistil. Moreover, disrupted auxin homeostasis in stamen filaments was associated with defective filament elongation. Our work reveals the key functions of ABCG1 and ABCG16 in reproductive development and provides clues for identifying ABCG1 and ABCG16 substrates in Arabidopsis.     

A role for the miR396/GRF network in specification of organ type during flower development,as supported by ectopic expression of Populus trichocarpa miR396c in transgenic tobacco

The MIR396 family, composed of ath‐miR396a and ath‐miR396b in Arabidopsis, is conserved among plant species and is known to target the Growth‐Regulating Factor (GRF) gene family. ath‐miR396 overexpressors or grf mutants are characterised by small and narrow leaves and show embryogenic defects such as cotyledon fusion. Heterologous expression of ath‐miR396a has been reported in tobacco and resulted in reduction of the expression of three NtGRF genes. In this study, the precursor of the Populus trichocarpa ptc‐miR396c, with a mature sequence identical to ath‐miR396b, was expressed under control of the CaMV35S promoter in tobacco. Typical phenotypes of GRF down‐regulation were observed, including cotyledon fusion and lack of shoot apical meristem (SAM). At later stage of growth, transgenic plants had delayed development and altered specification of organ type during flower development. The third and fourth whorls of floral organs were modified into stigmatoid anthers and fasciated carpels, respectively. Several NtGRF genes containing a miR396 binding site were found to be down‐regulated, and the cleavage of their corresponding mRNA at the miR396 binding site was confirmed for two of them using RACE‐PCR analysis. The data obtained agree with the functional conservation of the miR396 family in plants and suggest a role for the miR396/GRF network in determination of floral organ specification.     

Beautiful,complicated—and intelligent? Novel aspects of the thigmonastic stamen movement in Loasaceae

In a recent study we investigated the complex mechanisms regulating the pollen release via thigmonastic stamen movement found exclusively in Loasaceae subfamily Loasoideae. We demonstrated that stamen movement is modulated by abiotic (light and temperature) as well as biotic stimuli (pollinator availability and visitation frequency). This is explained as a mechanism to adjust the rate of stamen movement and thus pollen dispensation to different environmental conditions in order to optimize pollen transfer. Stamen movement is rapid and thus a near-immediate response to pollinator visits. However, Loasaceae flowers also show a response to biotic stimuli on a longer time scale, by adjusting the duration of both the staminate and the carpellate phase of the anthesis. We here present two additional data sets on species not previously studied, underscoring the shortening of the staminate phase in the presence of pollinator visits vs. their absence and the shortening of the carpellate phase after pollination. Overall, the plant shows not only a rapid but an “intelligent” reaction to its environment in adjusting anthesis and pollen presentation to a range of factors. The physiological and morphological bases of the stamen movement are poorly understood. Our previous study showed that there is no direct spatial relationship between the place of stimulation in the flower and the stamen bundle activated. We here further show the morphological basis for stamen movement from a reflexed into an erect position: Only the basal part of the filament curves around the receptacle, while the upper part of the filament retains its shape. We hypothesize that the stimulus is transmitted over the entire receptacle and the place of reaction is determined by stamen maturity, not the location of the stimulus.     

秦岭石蝴蝶人工种群花器变异现象研究

以人工栽培的秦岭石蝴蝶为实验材料,通过观察并记录花器官形态和数目的变化,初步探讨秦岭石蝴蝶花器变异规律,并分析了导致其变异的诱因。结果显示:(1)在观察的1 996朵秦岭石蝴蝶花朵中,发现了17种花冠变异类型、5种萼片变异类型和7种可育雄蕊变异类型,总变异率分别为34.57%、38.38%和32.67%。(2)秦岭石蝴蝶花梗或可分支,花梗苞片数目2～3枚。(3)相关性分析结果表明,下唇数目与可育雄蕊数目呈正相关,相关系数为0.927 4,而上唇数目与可育雄蕊数目呈负相关,相关系数为-0.481 1,结合花型图示分析这可能与秦岭石蝴蝶雄蕊着生于花冠下唇内侧近基部有关。该研究统计的秦岭石蝴蝶变异类型丰富,可能对于今后秦岭石蝴蝶的系统进化、花器官发育、生殖生态以及分子遗传方面研究奠定了基础,也为培育不同观赏价值的品种提供思路。     

Pollen,ovules, and pollination in pea: Success,failure, and resilience in heat

Field pea (Pisum sativum), a major grain legume crop, is autogamous and adapted to temperate climates. The objectives of this study were to investigate effects of high temperature stress on stamen chemical composition, anther dehiscence, pollen viability, pollen interactions with pistil and ovules, and ovule growth and viability. Two cultivars (“CDC Golden” and “CDC Sage”) were exposed to 24/18°C (day/night) continually or to 35/18°C for 4 or 7 days. Heat stress altered stamen chemical composition, with lipid composition of “CDC Sage” being more stable compared with “CDC Golden.” Heat stress reduced pollen viability and the proportion of ovules that received a pollen tube. After 4 days at 35°C, pollen viability in flower buds decreased in “CDC Golden,” but not in “CDC Sage.” After 7 days, partial to full failure of anthers to dehisce resulted in subnormal pollen loads on stigmas. Although growth (ovule size) of fertilized ovules was stimulated by 35°C, heat stress tended to decrease ovule viability. Pollen appears susceptible to stress, but not many grains are needed for successful fertilization. Ovule fertilization and embryos are less susceptible to heat, but further research is warranted to link the exact degree of resilience to stress intensity.     

Tomato stigma exsertion induced by high temperature is associated with the jasmonate signalling pathway

High temperature (HT) is becoming an increasingly serious factor in limiting crop production with global climate change. During hot seasons, owing to prevailing HT, cultivated tomatoes are prone to exhibiting stigma exsertion, which hampers pollination and causes fruit set failure. However, the underlying regulatory mechanisms of the HT‐induced stigma exsertion remain largely unknown. Here, we demonstrate that stigma exsertion induced by HT in cultivated tomato is caused by more seriously shortened stamens than pistils, which is different from the stigma exsertion observed in wild tomato species. Under the HT condition, the different responses of pectin, sugar, expansin, and cyclin cause cell wall remodelling and differentially localized cell division and selective cell enlargement, which further determine the lengths of stamens and pistils. In addition, auxin and jasmonate (JA) are implicated in regulating cell division and cell expansion in stamens and pistils, and exogenous JA instead of auxin treatment can effectively rescue tomato stigma exsertion through regulating the JA/COI1 signalling pathway. Our findings provide a better understanding of stigma exsertions under the HT condition in tomato and uncover a new function of JA in improving plant abiotic stress tolerance.     

Phylogeny of the core Malvales: evidence from ndhF sequence data

The monophyly of the group comprising the core malvalean families, Bombacaceae, Malvaceae, Sterculiaceae, and Tiliaceae, was recently confirmed by molecular studies, but the internal structure of this clade is poorly understood. In this study, we examined sequences of the chloroplast ndhF gene (aligned length 2226 bp) from 70 exemplars representing 35 of the 39 putative tribes of core Malvales. The monophyly of one traditional family, the Malvaceae, was supported in the trees resulting from these data, but the other three families, as traditionally circumscribed, are nonmonophyletic. In addition, the following relationships were well supported: (1) a clade, /Malvatheca, consisting of traditional Malvaceae and Bombacaceae (except some members of tribe Durioneae), plus Fremontodendron and Chiranthodendron, which are usually treated as Sterculiaceae; (2) a clade, /Malvadendrina, supported by a unique 21-bp (base pair) deletion and consisting of /Malvatheca, plus five additional subclades, including representatives of Sterculiaceae and Tiliaceae, and Durionieae; (3) a clade, /Byttneriina, with genera traditionally assigned to several tribes of Tiliaceae, plus exemplars of tribes Byttnerieae, Hermannieae, and Lasiopetaleae of Sterculiaceae. The most striking departures from traditional classifications are the following: Durio and relatives appear to be more closely related to Helicteres and Reevesia (Sterculiaceae) than to Bombacaceae; several genera traditionally considered as Bombacaceae (Camptostemon, Matisia, Phragmotheca, and Quararibea) or Sterculiaceae (Chiranthodendron and Fremontodendron) appear as sister lineages to the traditional Malvaceae; the traditional tribe Helictereae (Sterculiaceae) is polyphyletic; and Sterculiaceae and Tiliaceae, as traditionally circumscribed, represent polyphyletic groups that cannot sensibly be maintained with their traditional limits for purposes of classification. We discuss morphological characters and conclude that there has been extensive homoplasy in characters previously used to delineate major taxonomic groups in core Malvales. The topologies here also suggest that /Malvatheca do not have as a synapormophy monothecate anthers, as has been previously supposed but, instead, may be united by dithecate, transversely septate (polysporangiate) anthers, as found in basal members of both /Bombacoideae and /Malvoideae. Thus, “monothecate” anthers may have been derived at least twice, independently, within the /Bombacoideae (core Bombacaceae) and /Malvoideae (traditional Malvaceae).     

Incomplete feminisation by the microsporidian sex ratio distorter,Nosema granulosis,and reduced transmission and feminisation efficiency at low temperatures

We investigated the effects of temperature on transovarial transmission and feminisation by Nosema granulosis, a microsporidian sex ratio distorter of the brackish water amphipod Gammarus duebeni. There was no difference in parasite transmission efficiency to the F(1) eggs of infected females maintained under two temperature conditions, 5 and 10 degrees C (89 and 86%, respectively). When F(1) individuals were screened as adults, the proportion infected was also similar at both temperatures (74 and 75%, respectively). However, transmission to the eggs of the F(2) generation was significantly reduced at low temperatures (61% at 5 degrees C and 91% at 10 degrees C). In addition, feminisation efficiency was reduced substantially at low temperatures; at 10 degrees C, a calculated 85% of infected males were feminised, but at 5 degrees C only 49% were feminised. This is the first evidence for incomplete feminisation and temperature-dependent transmission and feminisation by this sex ratio distorter. We examine the consequences for parasite spread and maintenance in natural populations using a model to predict parasite prevalence in large populations. Reduced feminisation at low temperatures impedes the spread of the parasite so that it attains a substantially lower frequency, or may even be excluded, from host populations.