Heterogeneous expression patterns and separate roles of the SEPALLATA gene LEAFY HULL STERILE1 in grasses

SEPALLATA (SEP) genes exhibit distinct patterns of expression and function in the grass species rice (Oryza sativa) and maize (Zea mays), suggesting that the role of the genes has changed during the evolution of the family. Here, we examine expression of the SEP-like gene LEAFY HULL STERILE1 (LHS1) in phylogenetically disparate grasses, reconstruct the pattern of gene expression evolution within the family, and then use the expression patterns to test hypotheses of gene function. Our data support a general role for LHS1 in specifying determinacy of the spikelet meristem and also in determining the identity of lemmas and paleas; these two functions are separable, as is the role of the gene in specifying floret meristems. We find no evidence that LHS1 determines flower number; it is strongly expressed in all spikelet meristems even as they are producing flowers, and expression is not correlated with eventual flower number. LHS1 expression in only the upper flowers of the spikelet appears to be the ancestral state; expression in all flowers is derived in subfamily Pooideae. LHS1 expression in pistils, stamens, and lodicules varies among the cereals. We hypothesize that LHS1 may have affected morphological diversification of grass inflorescences by mediating the expression of different floral identity genes in different regions of the floret and spikelet.     

Cloning and characterization of a bamboo LEAFY HULL STERILE1 homologous gene.

A cDNA named DlMADS8 was isolated from the young spikelets of the sweet bamboo, Dendrocalamus latiflorus by rapid amplification of cDNA end (RACE). DNA sequence analysis showed that DlMADS8 was composed of full ORF and 3'UTR, but without 5'UTR. The cDNA contained 1059 nucleotides and encoded a putative protein of 244 amino acid residues. The gene displayed the structure of a typical plant MADS-box gene, which consisted of a MADS domain, K domain, a short I region, and the C-terminal region. Phylogenetic analysis of plant MADS-box genes based on amino acid sequences revealed that DlMADS8 was grouped into the AGAMOUS-LIKE 2 (AGL2)-like subfamily. It was homologous to the LEAFY HULL STERILE1 (LHS1) genes of grasses. To study the functions of it, DlMADS8 cDNA clone driven by the CaMV 35S promoter was transformed into Arabidopsis thaliana. Transgenic plants of DlMADS8 exhibited the phenotypes of curled leaves and early flowering. After bolting, three novel phenotypes related to inflorescence development were observed in different transgenic plants. No obvious homeotic conversions of floral organs were observed in all of the 35S::DllMADS8 transgenic Arabidopsis plants. These results indicated that DlMADS8 probably plays a role in floral meristem determinacy and is involved in controlling the flowering time of D. latiflorus.     

Genome organization and polyploid evolution in the genus Eleusine (Poaceae)

 Eleusine (Poaceae) includes six diploid and three polyploid species and has three basic chromosome numbers, x=8, 9 and 10. The species are annual as well as perennial and all are wild except E. coracana, which is cultivated for grain and fodder in Africa and the Indian subcontinent. Eleusine coracana and E. africana have the same genome and chromosome number (2n=36). Eleusine indica and E. floccifolia are identified as two genome donors to these polyploid species. Eleusine kigeziensis is the third polyploid species of the genus with 2n=38. Its genome may have come from E. jaegeri and from one of the species with x=9, most probably from E. indica. Eleusine indica, E. tristachya, E. floccifolia and E. intermedia with x=9 and two polyploid species, E. coracana and E. africana, are closely related and there is free genetic flow between them. Eleusine multiflora with x=8 is significantly different in morphology and at genomic level from other species. Eleusine jaegeri with x=10 is morphologically similar to E. indica, however, more information is needed to ascertain its position in the genus. Eleusine coracana, which is commonly called finger millet, is a potential and nutritious crop for the increasing population of the world, particularly in arid and semi-arid regions. It can also serve as a gene pool for various important characters and disease resistant genes. Received February 11, 2002; accepted May 27, 2002 Published online: October 14, 2002 Addresses of the authors: Madho Singh Bisht and Yasuhiko Mukai (e-mail: ymukai@cc.osaka-kyoiku.ac.jp), Laboratory of Plant Molecular Genetics, Division of Natural Science, Osaka Kyoiku University, 4-698-1 Asahigaoka, Kashiwara, Osaka 582-8582, Japan.     

Molecular evolution of FLORICAULA/LEAFY orthologs in the Andropogoneae (Poaceae)

Members of the grass family (Poaceae) exhibit a broad range of inflorescence structures and other morphologies, making the grasses an interesting model system for studying the evolution of development. Here we present an analysis of the molecular evolution of FLORICAULA/LEAFY-like genes, which are important developmental regulatory loci known to affect inflorescence development in a wide range of flowering plant species. We have focused on sequences from the Andropogoneae, a tribe within the grass family that includes maize (Zea mays ssp. mays) and Sorghum (Sorghum bicolor). The FLORICAULA/LEAFY gene phylogeny we generated largely agrees with previously published phylogenies for the Andropogoneae using other nuclear genes but is unique in that it includes both members of one of the many duplicate gene sets present in maize. The placement of these sequences in the phylogeny suggests that the duplication of the maize FLORICAULA/LEAFY orthologs, zfl1 and zfl2, is a consequence of a proposed tetraploidy event that occurred in the common ancestor of Zea and a closely related genus, Tripsacum. Our data are consistent with the hypothesis that the transcribed regions of the FLORICAULA/LEAFY-like genes in the Andropogoneae are functionally constrained at both nonsynonymous and synonymous sites and show no evidence of directional selection. We also examined conservation of short noncoding sequences in the first intron, which may play a role in gene regulation. Finally, we investigated the genetic diversity of one of the two maize FLORICAULA/LEAFY orthologs, zfl2, in maize and its wild ancestor, teosinte (Z. mays ssp. parviglumis), and found no evidence for selection pressure resulting from maize domestication within the zfl2-coding region.     

适应辐射类群穇属的系统学研究进展

综合花序拓扑学、比较形态学、分子系统发育、细胞遗传学等资料,对适应辐射类群穆属(Eleusine Gaertn.)的系统学研究进展进行了述评.穆属系统位置--Eleusiinae亚族成员得到分子系统发育证据的支持.该属具有3种花序类型、7个基因组类型、多倍体均由二倍体杂交起源、C4植物高度适应半湿润-半干旱镶嵌气候等特征.据可靠化石记载和现代地理分布推断,穆属很可能起源于东非,时间是晚中新世,而适应辐射则发生在上新世-中新世间隔.总的来说,分子系统发育、细胞遗传学、古地质、古气候数据的整合研究能够为穆属多倍体起源和谱系多样化历史提供令人信服的证据.     

穇属颖果微形态及其系统学意义

在电子扫描显微镜下观察了穆属(Eleusine Gaertn.)9种植物颖果的微形态特征.结果表明,穆属颖果有卵形、披针形和矩圆形3种腹面形状,压扁方式有背腹压扁及两侧压扁,腹面形态有平腹面和凹腹面两种,胚比为0.29～0.58,种脐比为0.10～0.24,颖果纹饰有2种,包括疣突状一级纹饰和疣突规则排列构成的复合网状...     

Ultrastructure and development of the megagametophyte in Eleusine tristachya (Lam.) Lam. (Poaceae)

Eleusine tristachya (Lam.) Lam. is native from subtropical South American climates. Widespread in Argentina and Uruguay, it is frequently found in landscape prairies of the province of Buenos Aires. Megasporogenesis and megagametogenesis in this species were studied using light and transmission microscopy. The ovule is hemitropous, bitegmic and tenuinucellate. The megaspore mother cell enlarges and undergoes meiosis division resulting in a T-shaped tetrad of megaspores. The three micropylar megaspores degenerate, and the chalazal one develops into the Polygonum-type megagametophyte. The synergid cells have the cytoplasm very electron dense because it has got a rich complement of organelles. The synergid wall is strongly thickened at the micropylar pole, developing the filiform apparatus. At maturity, the antipodals originate a wall with large projections into the cytoplasm, acquiring transfer cells characteristics. The antipodals cytoplasm, enriched with organelles shows a high metabolic activity, and it is suggested that these cells perform as an efficient system for metabolites transport.     

Allotetraploid origin and divergence in Eleusine (Chloridoideae, Poaceae): evidence from low-copy nuclear gene phylogenies and a plastid gene chronogram

Background and Aims Eleusine

(Poaceae) is a small genus of the subfamily Chloridoideae exhibiting considerable morphological and ecological diversity in East Africa and the Americas. The interspecific phylogenetic relationships of Eleusine are investigated in order to identify its allotetraploid origin, and a chronogram is estimated to infer temporal relationships between palaeoenvironment changes and divergence of Eleusine in East Africa.

Methods

Two low-copy nuclear (LCN) markers, Pepc4 and EF-1α, were analysed using parsimony, likelihood and Bayesian approaches. A chronogram of Eleusine was inferred from a combined data set of six plastid DNA markers (ndhA intron, ndhF, rps16-trnK, rps16 intron, rps3, and rpl32-trnL) using the Bayesian dating method.

Key Results

The monophyly of Eleusine is strongly supported by sequence data from two LCN markers. In the cpDNA phylogeny, three tetraploid species (E. africana, E. coracana and E. kigeziensis) share a common ancestor with the E. indica–E. tristachya clade, which is considered a source of maternal parents for allotetraploids. Two homoeologous loci are isolated from three tetraploid species in the Pepc4 phylogeny, and the maternal parents receive further support. The A-type EF-1α sequences possess three characters, i.e. a large number of variations of intron 2; clade E-A distantly diverged from clade E-B and other diploid species; and seven deletions in intron 2, implying a possible derivation through a gene duplication event. The crown age of Eleusine and the allotetraploid lineage are 3·89 million years ago (mya) and 1·40 mya, respectively.

Conclusions

The molecular data support independent allotetraploid origins for E. kigeziensis and the E. africana–E. coracana clade. Both events may have involved diploids E. indica and E. tristachya as the maternal parents, but the paternal parents remain unidentified. The habitat-specific hypothesis is proposed to explain the divergence of Eleusine and its allotetraploid lineage.     

Phylogeny of Eleusine (Poaceae: Chloridoideae) based on nuclear ITS and plastid trnT-trnF sequences

Phylogenetic relationships in the genus Eleusine (Poaceae: Chloridoideae) were investigated using nuclear ITS and plastid trnT-trnF sequences. Separate and combined data sets were analyzed using parsimony, distance, and likelihood based methods, including Bayesian. Data congruence was examined using character and topological measures. Significant data heterogeneity was detected, but there was little conflict in the topological substructure measures for triplets and quartets, and resolution and clade support increased in the combined analysis. Data incongruence may be a result of noise and insufficient information in the slower evolving trnT-trnF. Monophyly of Eleusine is strongly supported in all analyses, but basal relationships in the genus remain uncertain. There is good support for a CAIK clade (E. coracana subsp. coracana and africana, E. indica, and E. kigeziensis), with E. tristachya as its sister group. Two putative ITS homeologues (A and B loci) were identified in the allotetraploid E. coracana; the 'B' locus sequence type was not found in the remaining species. Eleusine coracana and its putative 'A' genome donor, the diploid E. indica, are confirmed close allies, but sequence data contradicts the hypothesis that E. floccifolia is its second genome donor. The 'B' genome donor remains unidentified and may be extinct.     

The dilemma of Guinea grass (Megathyrsus maximus): a valued pasture grass and a highly invasive species

On a global scale, invasive grasses threaten biodiversity and ecosystem function. Nevertheless, the importation of forage grasses is a significant economic force driven by globalization. Pastureland and rangeland are of critical economic and ecological importance, but novel grass species may lead to invasion. Recognizing that economically important species can also be ecologically damaging creates a contentious debate for land managers, policymakers, and ecologists alike. Many Afrotropical perennial grass species have been intentionally introduced pantropically given their high forage production and resistance to stress. However, these traits may also confer competitive ability, increasing the possibility of unintended escape and invasion. Further, these traits have posed challenges for traditional control methods using chemicals, prescribed fire, and mowing. The use of classic biological control may alleviate the ecological impact in invaded areas. In this literature synthesis we examine Guinea grass (Megathyrsus maximus); whose economic value in many countries is undeniable, yet its impact on native ecosystems is a mounting concern. First, we introduce Guinea grass taxonomy, general biology and ecology, and the geographic and genetic origins. Second, we review the economic value and the ecological impacts. Third, we review the control of Guinea grass in undesired areas using chemical and mechanical means. Finally, we review current efforts to use biological control.

     

Evidence from RAPD markers in the evolution ofEchinochloa millets (Poaceae)

Echinochloa (Poaceae) includes two domesticated species,Echinochloa utilis (Japanese barnyard millet) andE. frumentacea (Indian sawa millet) and 20–30 wild species. The two millets are morphologically very variable and overlap in spikelet and inflorescence characteristics. Both species are hexaploids based on x = 9. Cytogenetic studies point to the hexaploid wild speciesE. crusgalli andE. colona as possible progenitors ofE. utilis andE. frumentacea, respectively. The tetraploidE. oryzoides is considered as a possible genome donor to wild and domesticated barnyard millet. Markers from Random Amplified Polymorphic DNA method were used to assess the proposed phylogeny and examine the genetic diversity in both domesticated and wild species. The data were analyzed numerically.Echinochloa utilis andE. frumentacea appear very distinct, but grouped withE. crusgalli andE. colona, respectively. The tetraploidE. oryzoides show strong genetic affinity to theE. utilis—E. crusgalli group. The data are in general agreement with the cytogenetic information; however, some disagreements on the interpretation of some of the cytogenetic information is raised. The variability in DNA markers observed in the domesticated species, particularlyE. frumentacea, points to the feasibility of using RAPD markers in cultivar fingerprinting and breeding programs of these millets.     

Competition between Native Hawaiian Plants and the Invasive Grass Megathyrsus maximus: Implications of Functional Diversity for Ecological Restoration

Biodiversity loss is a global crisis, due primarily to habitat destruction and widespread nonnative invasions. Invasive grasses are particularly problematic in many tropical ecosystems, where they possess traits that promote their persistence and can drastically alter native plant communities. We explored the ecophysiological basis for restoring native Hawaiian dryland ecosystems currently dominated by the nonnative invasive grass Megathyrsus maximus (guinea grass) in a garden experiment. Three native species—Myoporum sandwicense (naio; canopy tree), Dodonaea viscosa (aalii; shrub), and Plumbago zeylanica (iliee; groundcover)—were grown with M. maximus at three levels of native functional diversity (one, two, or three species) while holding overall plant density constant. We tested which individual and functional combinations of native species were more productive and best suppressed M. maximus growth and reproduction. Megathyrsus maximus had 39–94% higher maximum photosynthetic rates (A_max) than native species and increasing native functional diversity did not affect M. maximus A_max. Aboveground, belowground, and total biomass of M. maximus varied with functional diversity, although intraspecific competition reduced growth as much as interspecific competition. Reproductive tiller production by M. maximus decreased significantly when planted with any of the native species and with increasing native functional diversity. These results indicate that high native functional diversity in an ecological restoration setting may aid in the control of a dominant invasive grass and the reintroduction of diverse native species. Recommendations for restoring degraded nonnative grasslands in Hawaii and throughout the tropics include selection of native species that are ecophysiologically competitive and have high functional diversity.     

Phylogeny and molecular evolution of the DMC1 gene within the StH genome species in Triticeae (Poaceae)

To estimate the phylogeny and molecular evolution of a single-copy nuclear disrupted meiotic cDNA (DMC1) gene within the StH genome species, two DMC1 homoeologous sequences were isolated from nearly all the sampled StH genome species and were analyzed with those from seven diploid taxa representing the St and H genomes in Triticeae. Sequence diversity patterns and genealogical analysis suggested that (1) there is a close relationship among North American StH genome species; (2) the DMC1 gene sequences of the StH genome species from North America and Eurasia are evolutionarily distinct; (3) the StH genome polyploids have higher levels of sequence diversity in the St genome homoeolog than the H genome homoeolog; (4) the DMC1 sequence may evolve faster in the polyploid species than in the diploids; (5) high dN and dN/dS values in the St genome within polyploid species could be caused by low selective constraints or AT-biased mutation pressure. Our result provides some insight on evolutionary dynamics of duplicate DMC1 gene, the polyploidization events and phylogeny of the StH genome species.     

Co-segregation of the gynomonoecious sex form1 gene (gsf1) of Tripsacum dactyloides (Poaceae) with molecular markers.

The only monogenic trait in Tripsacum to date was first identified in the prolific sex form variant Tripsacum dactyloides (L.) L. forma prolificum Dayton et Dewald. The expression of this trait is controlled by the presence of a single-gene, recessive pistillate mutation hereby designated the gynomonoecious sex form1 gene (gsf1), after the registered plant germplasm accession GSF-I (PI483447) from which it was first identified. This trait confers a high degree of feminization to the primarily male floral structure of the Tripsacum rachis. Two molecular markers were found to co-segregate with the gsf1 gene in a diploid (2n = 36) F2 population of Tripsacum dactyloides, where the female parent (GSF-I) had been previously determined to be homozygous recessive for the gene. Phenotypic scoring data were compared with restriction fragment length polymorphism data and linkage relationships were determined. The gsf1 gene is located ~7 cM from tda48, a Tripsacum-derived molecular marker, and ~9 cM from npi286, a maize-derived molecular marker. The marker npi286 also maps within ~5 cM of the tassel seed2 locus (ts2) of maize, which confers a similar change in the inflorescence of the maize tassel.     

Development of PCR primer sets for intron 1 of the low-copy gene LEAFY in Davalliaceae

? Premise of the study: Primers were designed for amplifying intron 1 of the single-copy nuclear LEAFY gene for species of Davalliaceae. ? Methods and Results: New primer sets were designed and successfully amplified for intron 1 of the LEAFY gene in 13 species representing the five genera of Davalliaceae. The orthology of these sequences was further confirmed by phylogenetic analyses. Site variation in LEAFY intron 1 sequences across genera of the Davalliaceae and among accessions of the Humata repens complex were 18% and 8%, respectively. Such variation was greater than that for the cpDNA atpB-rbcL intergenic spacer region across the same taxa and accessions. ? Conclusions: Using our newly designed primers, intron 1 of the LEAFY gene could be amplified for all species tested. In addition, this single-copy, biparentally inherited, and quickly evolving region showed considerable potential for addressing infraspecific-level questions.