The coexistence of binucleate and trinucleate pollen in Mitrephora macclurei Weerasooriya & R. M. K (Annonaceae)

Pollen of angiosperms may be either binucleate or trinucleate at maturity. According to Schürhoff–Brewbaker law, the binucleate state is pleisomorphic occuring in about 70% of all angiosperms, while the trinucleate state is advanced occurring in about 30% of the angiosperms. The derivation of trinucleate pollen from binucleate pollen is irreversible, thus trinucleate or mixed pollen was considered uncommon in early lineage. In the present work, using conventional paraffin section method, the coexistence of both binucleate and trinucleate pollen in the same intact anther were observed in Mitrephora macclurei, a species belonging to the eumagnoliid family Annonaceae. The increasing number of plants with trinucleate or mixed pollen discovered in early lineages, as well as climatic introduced changes in nuclear number challenge analyses of ancestral state and evolution of pollen nuclear number in angiosperms.     

Susceptibility of pollen to UV-B radiation: an assay of 34 taxa

Much of the ultraviolet-B radiation (UV-B) research on plants has concentrated on vegetative plant parts, and only a small fraction has dealt with the reproductive system. The present study analyzed pollen grains of 34 taxa germinated and grown under two levels of UV-B radiation (187 and 460 mW/m2) or no UV-B (control group). Visible radiation at 260 mmol/m/s was present in all treatments. Taxa included those with binucleate and trinucleate pollen types. We detected differences among species. A significant reduction in pollen germination occurred in only five species. Pollen tubes of >50% of the species showed significant reduction in length. Trinucleate pollen types were more likely to exhibit tube length reduction than the binucleate types. Proportionately more monocotyledonous species were sensitive to UV-B treatment than dicotyledonous species, and proportionately more wild species were sensitive than cultivated species and pollen collected from plants growing in the field were somewhat more sensitive than pollen collected from plants grown in the greenhouse. Species in which pollination occurred earlier in the season were more likely to be susceptible to UV-B radiation than those for which anthesis took place later in the season, suggesting a possible adaptation to UV-B radiation.     

细胞质雄性不育水稻幼穗和花粉发育期的蛋白酶活性变化

水稻细胞质雄性不育系珍汕97A,在花粉母细胞形成期以后可溶性蛋白含量迅速降低。单核期仅为花粉母细胞形成期的56％,二核期和三核期分别为花粉母细胞形成期的36．4％和30．3％。保持系珍汕97B虽在花粉母细胞形成期后可溶性蛋白含量降低,但在二核期则有所增高,表现为鞍形变化过程。在花粉母细胞形成期后,珍汕97A的酪蛋白水解酶活性增高,单核期为前一时期的1倍,而珍汕97B的酶活性增幅较小。第一次枝梗分化期珍汕97A的内肽酶活性较珍汕97B高,而雌雄蕊形成期两者的酶活皆降低。整个发育期中珍汕97A的内肽酶活性较珍汕97B高15％至55．5％。珍汕97A在幼穗第一次枝梗分化期时氨肽酶活性较珍汕97B高。第二次枝梗分化期后,不育系和保持系的氨肽酶活性均下降。但花粉母细胞形成期后,珍汕97A的氨肽酶活性迅速增高,三核期达最高水平,这可能意味着此时大量蛋白质被水解。蛋白酶类抑制剂的实验表明,在雌雄蕊形成期后不育系和保持系含有半胱氨酸型氨肽酶和约10％含金属型氨肽酶;而在三核期,保持系的氨肽酶则主要为半胱氨酸型。不育系三核期含有的BAPA内肽酶包括半胱氨酸型、丝氨酸型,以及部分含金属酶类;而保持系三核期的内肽酶仅为丝氨酸型。不育系肽酶类型的变化,可能反映酶基因编码的去抑制。     

RAPID SCREENING METHOD TO DETECT POTENTIAL BIPARENTAL INHERITANCE OF PLASTID DNA AND RESULTS FOR OVER 200 ANGIOSPERM SPECIES

We have developed a diagnostic method to screen rapidly for plant species potentially capable of biparental inheritance of plastid DNA using the DNA fluorochrome 4′,6-diamidino-2-phenylindole (DAPI) in conjunction with epifluorescence microscopy. Pollen shed from 235 plant species (including about 50 of agronomic importance) representing 80 families were screened. Putative plastid DNA was detected in the generative and/or sperm cells of pollen from 26 genera (43 species) representing 15 families. Plastid DNA was not detected in the generative or sperm cells of pollen from 192 plant species, thereby strongly suggesting that these species have only maternal inheritance. Our cytological diagnosis corroborated the known genetic evidence in 42 plant species and conflicted with the genetic reports in five species, which are discussed. The data suggest that biparental inheritance of plastids is rare; overall, it may occur in about 14% of flowering plant genera, examples of which are scattered among 19% of the families examined. This methodology also readily reveals whether pollen is bi- or trinucleate.     

木兰藤科系统位置评述

木兰藤科(Austrobaileyaceae)含1属2种,是系统学上最孤立的科之一。其花粉类似于最古老的被子植物化石之一:晚白垩世的棒纹粉。最新的分子系统发育研究结果表明,木兰藤科是现存被子植物的基部类群之一,其对于被子植物的起源与早期进化的研究具有重要价值。被子植物(有花植物)的起源和辐射一直是植物学家关注的热点。有关木兰藤科的系统位置一直存在争议。本文对该科系统位置的研究历史与现状进行评述。     

SYSTEMATIC SIGNIFICANCE OF POLLEN NUCLEAR NUMBER IN EUPHORBIACEAE,TRIBE EUPHORBIEAE

Pollen nuclear number is determined in 139 species of 5 genera in the Euphorbieae, subtribe Euphorbiinae. The 111 new determinations are tabulated along with previous reports, and the results indicate that the distribution of binucleate (II) and trinucleate (III) pollen is strongly associated with the taxonomic groupings within the Euphorbieae. Although binucleate pollen is probably primitive within the tribe Euphorbieae, as suggested by the nuclear condition in Neoguillauminia, the situation in Euphorbia still requires further elucidation. Within Euphorbia, the morphologically most primitive species studied have III pollen despite the fact that II pollen is presumably the original condition for the subtribe Euphorbiinae. In Euphorbia, II pollen only is reported from nine sections and III pollen only from ten sections, while in four sections (Esula, Goniostema, Aphyllis, and Deuterocalli) both II and III pollen have been found. The New World species of Euphorbia nearly all have III pollen, whereas the vast majority of the African succulents have II pollen. The genera of New World origin, Chamaesyce and Pedilanthus, have III pollen, while the African genera Monadenium and Synadenium have II pollen. Independent derivations of III pollen from II pollen appear to have occurred in sections Goniostema, Aphyllis, and Deuterocalli (all of subg. Euphorbia). There is no evidence that reversals from III to II pollen have occurred.     

Phylogenetic implications of pollen nuclear number in theAraceae

Data on pollen nuclear number are presented for 74 of the 111 currently accepted genera of theAraceae — one of only three monocot families in which both bi- and trinucleate pollen are known to occur. Binucleate pollen, which characterizes 65% of aroid genera, is regarded as the primitive type inAraceae and monocots as a whole. The trend toward the trinucleate condition appears to be irreversible, and to have occurred many times within the family.Syngonium becomes the sixth angiosperm genus in which both character-states are known to occur; otherwise, the character is highly conservative at the generic level. Pollen nuclear number shows no evident correlation with pollen size, starch content or pollinator type.     

DNA methylation analysis of a male reproductive organ specific gene (MROS1) during pollen development.

Pollen grains of angiosperm plants represent a good model system for studies of chromatin structure and remodelling factors, but very little is known about the DNA methylation status of particular genes in pollen. In this study, we present an analysis of the DNA methylation patterns of the MROS1 gene, which is expressed in the late phases of pollen development in Silene latifolia (syn. Meladrium album). The genomic sequencing technique revealed similar DNA methylation patterns in leaves, binucleate pollen, and trinucleate pollen. Extremely high DNA methylation levels occurred in the CG dinucleotides of the upstream region (99%), whereas only a low level of CG methylation was observed in the transcribed sequence (7%). Low levels of methylation were also observed in asymmetric sequences (in both regions; 2% methylated). The results obtained in the MROS1 gene are discussed in consequence with the immunohistochemical data showing a hypermethylation of DNA in the vegetative nucleus.     

湖北光周期敏感核不育水稻农垦58S幼穗和花药发育期的蛋白酶活性变化

湖北光周期敏感核不育水稻农垦585的可育与不育株在一次枝梗期的幼穗中可溶性蛋白含量相近。雌雄蕊形成期和花粉母细胞分化期的可有株可溶性蛋白较不育株的分别高21％和12．3％;可有株在花粉二核和三核期亦有较高含量。不育株在雌雄蕊形成期后,酪蛋白酶活性迅速增高,而可有株至二核期才增高。不育株除在二次枝梗期的内肽酶活性略低外,其余阶段酶的活性较高。育性转变时内肽酶类型亦发生变化。可有株有较高的氨肽酶活性．     

Self-sterility in flowering plants: preventing self-fertilization increases family diversification rates

Background and Scope

New data are presented on the distribution and frequency of self-sterility (SS) – predominantly pre-zygotic self-incompatibility (SI) systems – in flowering plants and the hypothesis is tested that families with self-sterile taxa have higher net diversification rates (DRs) than those with exclusively self-compatible taxa using both absolute and relative rate tests.

Key Results

Three major forms of SI systems (where pollen is rejected at the stigmatic, stylar or ovarian interface) are found to occur in the oldest families of flowering plants, with times of divergence >100 million years before the present (mybp), while post-fertilization SS and heterostyly appear in families with crown ages of 81 and 87 mybp, respectively. It is also founnd that many (22) angiosperm families exhibit >1 SI phenotype and that the distribution of different types of SS does not show strong phylogenetic clustering, collectively suggesting that SS and SI systems have evolved repeatedly de novo in angiosperm history. Families bearing self-sterile taxa have higher absolute DRs using all available calibrations of the angiosperm tree, and this affect is caused mostly by the high DR of families with homomorphic SI systems (in particular stigmatic SI) or those in which multiple SS/SI phenotypes have been observed (polymorphic). Lastly, using sister comparisons, it is further demonstrated that in 29 of 38 sister pairs (including 95 families), the self-sterile sister group had higher species richness and DR than its self-compatible sister based on either the total number of taxa in the clade with SS or only the estimated fraction to harbour SS based on literature surveys.

Conclusions

Collectively, these analyses point to the importance of SS, particularly pre-zygotic SI in the evolution of flowering plants.     

The identification of fossil angiosperm pollen and its bearing on the time and place of the origin of angiosperms

Studies in the 1970's reporting the occurrence of fossil pollen types in the Cretaceous, coupled with surveys of extant pollen morphology of primitive flowering plants, laid the foundation for proposing a Lower Cretaceous origin of angiosperms. Over the last 30 years, morphological, ultrastructural, and ontogenetic studies of both extant and fossil pollen have provided an array of new characters, as well as greater resolution in defining character polarities. Moreover, a range of fossil pollen types exhibiting angiosperm characters occur in low frequency within Triassic and Jurassic sediments. The pollen data provide evidence of a pre-Cretaceous origin of angiosperms. Speciation and extinction rates were likely equal during the Triassic and Jurassic, resulting in the paucity of angiosperm pollen types from different geographic areas in the Atlantic – South American/African rift zone. It was not until the Lower Cretaceous that origination rates exceed extinction rates, resulting in the subsequent diversification of angiosperms and the origin of the eudicots.     

Convergent evolution of seed dispersal by ants,and phylogeny and biogeography in flowering plants: A global survey

Seed dispersal is a fundamental life history trait in plants. Although the recent surge of interest in seed dispersal by ants (myrmecochory) has added greatly to knowledge on the ecology of seed dispersal and ant–plant mutualisms, myrmecochory also represents a unique opportunity to examine the links between seed dispersal and evolution in flowering plants. Here we review the taxonomic, phylogenetic and biogeographic distribution of myrmecochory in flowering plants. Myrmecochory is mediated by elaiosomes, i.e., lipid-rich seed appendages that attract ants and serve as rewards for dispersal. We surveyed the literature for evidence of elaiosomes in angiosperm plants to estimate the global prevalence of myrmecochory. We then searched the literature for phylogenetic reconstructions to identify myrmecochorous lineages and to estimate the minimum number of independent evolutionary origins of myrmecochory. We found that myrmecochory is present in at least 11 000 species or 4.5% of all species, in 334 genera or 2.5% of all genera and in 77 families or 17% of all families of angiosperm plants. We identified at least 101, but possibly up to 147, independent origins of myrmecochory. We estimated three or more origins in 13 families and found that at least half the genera are myrmecochorous in 10 families. Most myrmecochorous lineages were Australian, South African or northern temperate (Holarctic). A mapping of families containing myrmecochorous genera on a dated angiosperm supertree showed that myrmecochory has evolved in most of the major angiosperm lineages and that it is more frequent in younger families (crown group age <80 million years) than in older ones. We suggest that the relatively low physiological and energetic costs of producing an elaiosome and the consistent selective benefits of myrmecochory (dispersal, protection from seed predators and fire, safe and nutrient-rich microsites) explain the numerous evolutionary and developmental origins of myrmecochory in angiosperm plants, and we propose that elaiosomes thus provide one of the most dramatic examples of convergent evolution in biology.     

Respiration and Vitality of Binucleate and Trinucleate Pollen

The respiration and vitality of ungerminated bi- and trinucleate pollen were studied in order to determine the influence of relative humidity and temperature on metabolic activity. The gas exchange, germination capacity and staining with tetrazolium bromide were followed under standardized conditions. A constant respiration rate occurred under conditions of high relative humidity (97%). Per mg pollen, the trinucleate grains of Compositae and Gramineae respired 2 to 3 times as intense as 6 species of binucleate grains. Per unit of pollen protein the differences were even larger. In contrast to binucleate pollen, the longevity of trinucleate pollen was very short and the ability to germinate was lost twice as fast as the respiration capacity. This limits the use of tetrazolium bromide as an indicator of viability. At reduced relative humidities respiration was strongly restricted, but the longevity of bi- and trinucleate pollen considerably increased. Pollen of Gramineae, however, was very sensitive to changes in relative humidity; short exposure to low relative humidity decreased both the vitality and the capacity to respire.     

Reproductive ecology and pollen representation among neotropical trees

Three years of pollen trapping data from Barro Colorado Island, Panama, are compared with local vegetation inventories. Two hypotheses relating pollen representation to ‘messy’ pollination and flower form are tested. Dioecious taxa were found to be over‐represented in pollen spectra compared with their occurrence in local forests. Similarly, anemophilous and ‘messy’ pollination types were found to be over‐represented. While anemophilous taxa were the best dispersed pollen types, zoophilous taxa were also well‐represented in dispersal classes of 20–40 m and > 40 m. Thus pollen arriving to lake sediments is most likely to be from anemophilous species or those zoophilous species exhibiting ‘messy’ pollination syndromes. Pollination mechanisms will predictably bias the fossil record against certain flower morphologies. These data are of significance to those using the fossil pollen record to reconstruct the timing and sequence of angiosperm evolution. These data help prioritize plants to be included in modern pollen reference collections and to focus the search for ‘unknown’ types on most‐likely candidate families.     

Pollen development in Annona cherimola Mill. (Annonaceae). Implications for the evolution of aggregated pollen

Background  

In most flowering plants, pollen is dispersed as monads. However, aggregated pollen shedding in groups of four or more pollen grains has arisen independently several times during angiosperm evolution. The reasons behind this phenomenon are largely unknown. In this study, we followed pollen development in Annona cherimola, a basal angiosperm species that releases pollen in groups of four, to investigate how pollen ontogeny may explain the rise and establishment of this character. We followed pollen development using immunolocalization and cytochemical characterization of changes occurring from anther differentiation to pollen dehiscence.     

Taxonomic description of in situ bee pollen from the middle Eocene of Germany

The middle Eocene Messel and Eckfeld localities are renowned for their excellently preserved faunas and diverse floras. Here we describe for the first time pollen from insect-pollinated plants found in situ on well-preserved ancient bees using light and scanning electron microscopy. There have been 140 pollen types reported from Messel and 162 pollen types from Eckfeld. Here we document 23 pollen types, six from Messel and 18 from Eckfeld (one is shared). The taxa reported here are all pollinated by insects and mostly not recovered in the previously studied dispersed fossil pollen records. Typically, a single or two pollen types are found on each fossil bee specimen, the maximum number of distinct pollen types on a single individual is five. Only five of the 23 pollen types obtained are angiosperms of unknown affinity, the remainder cover a broad taxonomic range of angiosperm trees and include members of several major clades: monocots (1 pollen type), fabids (7), malvids (4), asterids (5) and other core eudicots (1). Seven types each can be assigned to individual genera or infrafamilial clades. Since bees visit only flowers in the relative vicinity of their habitat, the recovered pollen provides a unique insight into the autochthonous palaeo-flora. The coexistence of taxa such as Decodon, Elaeocarpus, Mortoniodendron and other Tilioideae, Mastixoideae, Olax, Pouteria and Nyssa confirms current views that diverse, thermophilic forests thrived at the Messel and Eckfeld localities, probably under a warm subtropical, fully humid climate. Our study calls for increased attention to pollen found in situ on pollen-harvesting insects such as bees, which can provide new insights on insect-pollinated plants and complement even detailed palaeo-palynological knowledge obtained mostly from pollen of wind-pollinated plants in the dispersed pollen record of sediments. In the case of Elaeocarpus, Mortoniodendron, Olax and Pouteria the pollen collected by the middle Eocene bees represent the earliest unambiguous records of their respective genera.     

Protein synthesis of binucleate and trinucleate pollen and its relationship to tube emergence and growth

Under humid conditions, both bi- and trinucleate pollen species incorporate, on the average, very low amounts of leucine, e.g., 0.4 pmol min^-1mg pollen^-1. During germination in vitro, however, the two types of pollens greatly differ in their capacity for protein synthesis.Binucleate pollen species such as Typha, which are characterized by slow respiration in humid air and prolonged lag periods during germination in vitro, contain large amounts of monoribosomes at dehiscence. Polyribosomes are formed soon after the pollen is wetted in the germination medium, and a considerable incorporation of leucine is initiated after 10–15 min. More rapidly respiring, binucleate pollen showing a short lag period, such as Tradescantia, may already contain many polysomes at dehiscence and incorporate leucine within 2 min of germination. However, rapidly respiring, trinucleate Compositae pollen contains very limited amounts of ribosomal material and never attains any substantial level of incorporation. Cycloheximide completely inhibits both protein synthesis and tube emergence and growth in the slowly respiring, binucleate pollen species. The more rapidly respiring types are less dependent on protein synthesis, while germination of the phylogenetically advanced, trinucleate Compositae pollen proceeds completely independently. It is concluded that the level of phylogenetic advancement of the male gametophyte is characterized by its overall state of metabolic development at dehiscence rather than by the number of its generative cells.Abbreviations BSA bovine serum albumin - CHI cycloheximide - EGTA ethyleneglycol-bis(-aminoethyl ether) N,N-tetraacetic acid - RH relative humidity - TCA trichloroacetic acid     

K型温敏雄性不育系KTP116A育性转换规律的研究

基于两系杂交小麦系统的K型（Aegilops kotshyi）细胞质小麦温敏雄性不育系,在中国北方小麦正常生长季节表现完全雄性不育,可用于杂交小麦种子生产,在反季节播种表现育性部分恢复,是一类非常有利用价值的小麦温敏雄性不育系。本研究以小麦温敏雄性不育系KTP116A为材料,采用人工气候箱控温和涂抹法压片的方法对其育性转换的关键时期及败育的花粉发育过程进行更为精细的研究。研究发现,KTP116A可育环境下能正常进行减数分裂并发育为成熟花粉粒;不育环境下只有少数能经过第二次有丝分裂产生两个精细胞,大部分为二核期花粉粒、异常花粉粒和没有原生质体的空壳花药不开裂,自然条件下不散粉;KTP116A育性敏感期为两个时期,减数分裂期至单核早期和二核晚期至三核期,在自然条件下温度感应历期为10d左右;从小麦外部发育进程来看,抽穗前的2-8d和扬花前的3-5d为其敏感时期。这对于进一步研究败育的机理,分析败育的原因和更好的利用温敏不育系奠定基础。     

Substrate-dependent modulation of ATPase activity by Na+ and K+ in roots of Plantago species

The equal rates of water vapour absorption by both bi- and trinucleate pollen indicate that their widely-differing rates of respiration have an intrinsic, biochemical basis. This was investigated with various metabolic inhibitors that were previously introduced into dry pollen via anhydrous acetone. The uncoupler, carbonyl cyanide m-chlorophenyl hydrazone, inhibited the O₂ uptake of rapidly respiring pollen and stimulated that of slowly respiring types to similar absolute values, that probably reflect the rates of substrate transport across the mitochondrial membranes. The extent of inhibition of the O₂ uptake by oligomycin, dicyclohexyl carbodiimide, antimycin A, and salicyl hydroxamic acid, alone and in combinations, indicates that hardly any oxidative phosphorylation and anabolic activities occur in slowly respiring, binucleate pollen species, having low-developed mitochondria and high energy charge values. The presence of the alternative pathway was insignificant. In other binucleate pollen species, characterized by recognizable mitochondria and low energy charge values, a limited ATP synthesis was established. The low energy charge values point to imbalance between phosphorylative and anabolic activities. In rapidly respiring, trinucleate pollen, containing well-developed mitochondria, a significant activity of the alternative oxidase was found. The energy charge values were high notwithstanding the large demand for ATP, mounting to 1.7 μmol h^?1 (mg pollen)^?1. In some pollen species, oligomycin highly stimulated the flow of electrons through the cytochrome pathway, which made an estimation of the ATP synthesis impossible.     

Genetic diversity,distributional barriers and rafting continents — more thoughts on the evolution of mangroves

Without continental drift, the diversity and distribution of many species, including mangrove plants, would be very different today. First, there would be fewer pantropic genera and many more endemics. Second, their characteristics would not be as common and widespread as some are today. Continental drift has brought about the massive mixing and dispersal of genes in geologically recent times, greatly enhancing the evolutionary process; particularly for flowering plants — the angiosperms, which evolved during the period.Mangrove plants are comprised of approximately 70 species from 20 quite different angiosperm families. Most taxa are characterized by special physiological abilities and structural forms, enabling them to live in both seasonally fluctuating saline conditions, and water-saturated soils. Their occurrence is mostly tropical, perhaps because of harsh physiological conditions of intertidal habitats; but distributions of specific taxa do not fully concur with the idea of a completely tropical evolution, at least for some important species.At least one genus of mangrove tree, Avicennia, occurs around the world, chiefly in tropical estuarine habitats, although they also range into temperate latitudes, especially in the south. Around the world, there are no more than ten species of Avicennia recognised today, but their diagnostic determinants were inadequate prior to recent studies using both numerical analyses of morphological parameters and isozymes. Such analyses significantly reduced the number of apparent species, notably around Australia, and provided a basis for the revision of distributional records throughout the Indo West Pacific region. One species, A. marina, was found to be widespread and morphologically variable with genes divided into characteristic groupings of at least three geographic areas in the region. Based on these findings, there are several novel inferences to be made regarding the evolution of this genus. A western Gondwanan origin is proposed, with subsequent radiation of several taxa facilitated via the tectonic dispersal of southern continental fragments.