Evolution of Oleosin in Land Plants

Oleosins form a steric barrier surface on lipid droplets in cytoplasm, preventing them from contacting and coalescing with adjacent droplets. Oleosin genes have been detected in numerous plant species. However, the presence of oleosin genes in the most basally diverging lineage of land plants, liverworts, has not been reported previously. Thus we explored whether liverworts have an oleosin gene. In Marchantia polymorpha L., a thalloid liverwort, one predicted sequence was found that could encode oleosin, possessing the hallmark of oleosin, a proline knot (-PX₅SPX₃P-) motif. The phylogeny of the oleosin gene family in land plants was reconstructed based on both nucleotide and amino acid sequences of oleosins, from 31 representative species covering almost all the main lineages of land plants. Based on our phylogenetic trees, oleosin genes were classified into three groups: M-oleosins (defined here as a novel group distinct from the two previously known groups), low molecular weight isoform (L-oleosin), and high molecular weight isoform (H-oleosin), according to their amino-acid organization, phylogenetic relationships, expression tissues, and immunological characteristics. In liverworts, mosses, lycophytes, and gymnosperms, only M-oleosins have been described. In angiosperms, however, while this isoform remains and is highly expressed in the gametophyte pollen tube, two other isoforms also occur, L-oleosins and H-oleosins. Phylogenetic analyses suggest that the M-oleosin isoform is the precursor to the ancestor of L-oleosins and H-oleosins. The later two isoforms evolved by successive gene duplications in ancestral angiosperms. At the genomic level, most oleosins possess no introns. If introns are present, in both the L-isoform and the M-isoform a single intron inserts behind the central region, while in the H-isoform, a single intron is located at the 5′-terminus. This study fills a major gap in understanding functional gene evolution of oleosin in land plants, shedding new light on evolutionary transitions of lipid storage strategies.     

Evolution of Reproductive Organs in Land Plants

LEAFY gene is the positive regulator of the MADS-box genes in flower primordia. The number of MADS-box genes presumably increased by gene duplications before the divergence of ferns and seed plants. Most MADS-box genes in ferns are expressed similarly in both vegetative and reproductive organs, while in gymnosperms, some MADS-box genes are specifically expressed in reproductive organs. This suggests that (1) the increase in the number of MADS-box genes and (2) the subsequent recruitment of some MADS-box genes as homeotic selector genes were important for the evolution of complex reproductive organs. The phylogenetic tree including both angiosperm and gymnosperm MADS-box genes indicates the loss of the A-function genes in the gymnosperm lineage, which is presumably related to the absence of perianths in extant gymnosperms. Comparison of expression patterns of orthologous MADS-box genes in angiosperms, Gnetales, and conifers supports the sister relationship of Gnetales and conifers over that of Gnetales and angiosperms predicted by phylogenetic trees based on amino acid and nucleotide sequences. Received 30 July 1999/ Accepted in revised form 9 September 1999     

MIR166基因家族在陆生植物中的进化模式分析

MicroRNA（miRNA）是一类广泛存在于真核生物中的具有转录后水平调控功能的内源非编码小分子RNA。在植物中．miRNA通过对靶基因的剪切或沉默来实现对植物生命活动的调控,它是基因表达调控网络的重要组成部分。miR165／166（miR166）是陆生植物中最为古老的MIRNA家族之一,它通过对3型同源异域型-亮氨酸拉链（1id—ZIPⅢ）等靶标的调控,在植物的众多发育时期起着关键的调控作用。本文分析了MIR166基因在陆生植物中的进化关系,并对MIR166在基部陆生植物小立碗藓（Physcomitrella patens）中的复制及进化进行了研究。此外,HD—ZIPⅢ蛋白是植物中重要的一类转录因子,miR166对HD-ZIP Ⅲ基因的调控作用在陆地植物保守的存在,本文对HD—ZIP Ⅲ基因和miR166在进化中的相互作用进行了初步的探讨。     

Primary Cell Wall Structure in the Evolution of Land Plants

Investigation of the primary cell walls of lower plants improves our understanding of the cell biology of these organisms but also has the potential to improve our understanding of cell wall structure and function in angiosperms that evolved from lower plants. Cell walls were prepared from eight species, ranging from a moss to advanced gymnosperms, and subjected to sequential chemical extraction to separate the main polysaccharide fractions. The glycosyl compositions of these fractions were then determined by gas chromatography. The results were compared among the eight plants and among data from related studies reported in the existing published reports to identify structural features that have been either highly conserved or clearly modified during evolution. Among the highly conserved features are the presence of a cellulose framework, the presence of certain hemicelluloses such as xyloglucan, and the presence of rhamnogalacturonan II, a domain in pectic polysaccharides. Among the modified features are the abundance of mannosyl-containing hemicelluloses and the presence of methylated sugars.     

Generative Cell Specification Requires Transcription Factors Evolutionarily Conserved in Land Plants

1. Download high-res image (204KB)
2. Download full-size image

     

An Evolutionarily Conserved Signaling Mechanism Mediates Far-Red Light Responses in Land Plants

Phytochromes are plant photoreceptors important for development and adaptation to the environment. Phytochrome A (PHYA) is essential for the far-red (FR) high-irradiance responses (HIRs), which are of particular ecological relevance as they enable plants to establish under shade conditions. PHYA and HIRs have been considered unique to seed plants because the divergence of seed plants and cryptogams (e.g., ferns and mosses) preceded the evolution of PHYA. Seed plant phytochromes translocate into the nucleus and regulate gene expression. By contrast, there has been little evidence of a nuclear localization and function of cryptogam phytochromes. Here, we identified responses to FR light in cryptogams, which are highly reminiscent of PHYA signaling in seed plants. In the moss Physcomitrella patens and the fern Adiantum capillus-veneris, phytochromes accumulate in the nucleus in response to light. Although P. patens phytochromes evolved independently of PHYA, we have found that one clade of P. patens phytochromes exhibits the molecular properties of PHYA. We suggest that HIR-like responses had evolved in the last common ancestor of modern seed plants and cryptogams and that HIR signaling is more ancient than PHYA. Thus, other phytochromes in seed plants may have lost the capacity to mediate HIRs during evolution, rather than that PHYA acquired it.     

Motile Gametes of Land Plants: Diversity,Development, and Evolution

Referee: Professor Jeffrey Duckett, School of Biological Sciences, Queen Mary and Westfield College, University of London, Mile End Road, London, E1 4NS, UK Spermatogenesis is a morphogenetic system in plants that is unparalled in its potential to yield diverse and informative structural and developmental data. The unquestionable homology of terrestrial plant spermatozoids to each other and to gametes of related lineages allows an examination of cellular evolution and provides sound data for phylogenetic analyses. In this review we examine the architecture and ontogeny of motile male gametes among major groups of land plants. We begin with a historical perspective that emphasizes the utility of spermatogenesis in understanding cellular evolution and in determining phylogenetic relationships. A cladistic analysis of data based solely on spermatogenesis and a conceptual phylogeny based on combined morphological and molecular data serve as the basis for the comprehensive discussion of architectural and developmental features of plant spermatozoids. Spermatozoids of green plants have two fundamental architectural designs: biflagellated or multiflagellated. Biflagellated gametes vary among basal archegoniates and charophytes in degree of coiling, position, and substructure of the basal bodies and number of organelles. Hornwort spermatozoids are simple, bilaterally symmetrical, and uniquely exhibit a right-handed coil. An autapomorphy among setaphytes (a clade containing mosses and liverworts) is the production of coiled biflagellated sperm cells with dimorphic staggered basal bodies. Like bryophytes, gametes of most lycophytes are biflagellated; two exceptions are Isoëtes and Phylloglossum, taxa that independently evolved multiflagellated sperm cells with approximately 20 flagella. Developmental information, especially related to the origin and development of the locomotory apparatus, are essential to determine structural homology among these taxa. Evaluation of the more complicated multiflagellated gametes of other vascular plants reveals similarities that support a monophyletic fern, Equisetum and Psilotum assemblage. Autapomorphies of this clade include the arrangement of the microtubular cytoskeleton, origin of the locomotory apparatus, and structural details of the basal bodies and multilayered structure. Sperm cell development in archegoniates involves the complete transformation of virtually every cellular component. Crucial to this process are proteinaceous elements of the cytoskeleton. Complex microtubule arrays unique to these cells include the spline, basal bodies, and flagella. The discrete microtubule-organizing centers (MTOCs) that generate these cytoskeletal arrays are equally complex and enable the examination of molecular constituents and ontogenetic modifications. The protein centrin is found in a variety of structures, including the diverse MTOCs and the locomotory apparatus. Actin plays a role in organellar shaping and positioning as well as in cytoplasmic deletion and the maintenance of spatial integrity in the mature cell. We conclude with an overview of the current and potential utility of male gametogenesis as an informative system in approaching fundamental questions relating to cellular differentiation and motility. Characterization of motility mutants will elucidate genetic control of structure-function relationships among cellular components, while biochemical and molecular investigations provide crucial data on the mechanism of development. The examination of spermatogenesis in additional taxa is essential to characterize further developmental variations. Moreover, such studies provide a more comprehensive understanding of plant biodiversity at the cellular level and lead to even greater phylogenetic resolution from this elegant morphogenetic system.