一个水稻内颖退化突变体的形态特征及基因的精确定位

水稻产量和品质受花器官发育的直接影响,因此对水稻颖花发育机理的研究将有助于水稻产量提高和品质的改良。文章利用60Coγ射线辐照亲本8PW33(籼稻背景)获得一个性状能稳定遗传的内颖退化突变体(编号:MU102),并对其农艺性状和花器官进行了观察和分析。结果显示,相对于野生型,该突变体的株高、每穗总粒数及剑叶宽均显著增加,而结实率则显著降低,差异均达显著水平。解剖镜下观察表明,该突变体内颖退化,外颖弯曲呈现镰刀状,其余器官与野生型表型基本一致。扫描电镜观察显示,突变体与野生型叶片维管束的结构组成以及外颖表皮细胞组成、排列均正常,没有明显差异;与野生型相比,突变体内颖表皮细胞排列较为紧密,推测可能是内颖收缩退化导致的。遗传分析显示该突变性状是由隐性单基因控制,并命名为pd2。利用实验室现有的SSR分子标记将PD2基因定位于水稻第9号染色体上,通过进一步扩大群体和开发新的Indel标记,将PD2基因定位在2个Indel标记之间,两者间的物理距离大约是82 kb。在该物理区间内有一个已经克隆的内颖发育基因REP1,经过测序和比对分析,推测REP1与PD2为等位基因。     

ELE restrains empty glumes from developing into lemmas

Although there is evident homology among reproductive organs when comparing Poaceae(grass)and eudicots,the identity of grass specific organs,such as lodicules,palea,lemma,and glumes has been the subject of a vast and largely inconclusive discussion.Here we provide some direct evidence to support the idea that the empty glumes of rice(Oryza sativa)are counterparts of lemmas.We show that the development of empty glumes is regulated by ELE(elongated empty glume),which belongs to a plant specific novel gene family.Mutations at the ELE locus cause elongated empty glumes,which mimic the lemmas and have the epidermal morphology of lemmas with four or five vascular bundles.As a nuclear-localized gene,ELE is specifically expressed at the empty glumes of immature spikelets,and its ectopic expression causes many floral development defects,including lemma-like palea,extra palea-like structures,elongated lodicules,extra stamens and stigmas.Our result suggests that empty glumes are lemmas of the sterile florets located at the lateral side of the rice spikelet,and ELE acts as a regulator restraining its growth to maintain its small size in wild-type plants.     

Evidence for distinct roles of the SEPALLATA gene LEAFY HULL STERILE1 in Eleusine indica and Megathyrsus maximus (Poaceae)

LEAFY HULL STERILE1 (LHS1) is an MIKC-type MADS-box gene in the SEPALLATA class. Expression patterns of LHS1 homologs vary among species of grasses, and may be involved in determining palea and lemma morphology, specifying the terminal floret of the spikelet, and sex determination. Here we present LHS1 expression data from Eleusine indica (subfamily Chloridoideae) and Megathyrsus maximus (subfamily Panicoideae) to provide further insights into the hypothesized roles of the gene. E. indica has spikelets with three to eight florets that mature acropetally; E. indica LHS1 (EiLHS1) is expressed in the palea and lemma of all florets. In contrast, M. maximus has spikelets with two florets that mature basipetally; M. maximus LHS1 (MmLHS1) is expressed in the palea and lemma of the distal floret only. These data are consistent with the hypothesis that LHS1 plays a role in determining palea and lemma morphology and specifies the terminal floret of basipetally maturing grass spikelets. However, LHS1 expression does not correlate with floret sex expression; MmLHS1 is restricted to the bisexual distal floret, whereas EiLHS1 is expressed in both sterile and bisexual floret meristems. Phylogenetic analyses reconstruct a complex pattern of LHS1 expression evolution in grasses. LHS1 expression within the gynoecium has apparently been lost twice, once before diversification of a major clade within tribe Paniceae, and once in subfamily Chloridoideae. These data suggest that LHS1 has multiple roles during spikelet development and may have played a role in the diversification of spikelet morphology.     

Significance of consensus CYC-binding sites found in the promoters of both ChCYC and ChRAD genes in Chirita heterotricha (Gesneriaceae)

CYC-like genes are widely conserved in controlling floral dorsoventral asymmetry (zygomorphy) through persistent expression in corresponding domains in core eudicots. To understand how CYC-like gene expression is maintained during flower development, we selected Chirita heterotricha as a material and isolated the promoter sequences of the ChCYCIC and ChCYCID genes, homologs of CYC, by inverse polymerase chain reaction. Further promoter analyses led to the identification of a putative cis-regulatory element in each promoter matching the consensus DNA binding site for Antirrhinum CYC protein: GGCCCCTC at-165 for ChCYC1C, and GGCCCCCC at-163 for ChCYCID. This indicates that both the ChCYCIC and ChCYC1D genes have probably evolved autoregulatory loops to sustain their expression in developing flowers. We also isolated the coding and promoter sequences of the ChRAD gene, a homolog of Antirrhinum RAD. Promoter analysis showed that the ChRAD gene promoter also contained a putative CYC-binding site (GGCCCAC at -134). Therefore, ChRAD is likely a direct target of the ChCYC1 genes, which is similar to Antirrhinum RAD. These results imply that the establishment of floral zygomorphy in Chirita may have been achieved by the evolution of an autoregulatory loop for CYC-like genes,which was probably accompanied by simultaneous co-option of the RAD-like gene into their regulatory network.     

Molecular and genetic analyses of the silky1 gene reveal conservation in floral organ specification between eudicots and monocots

The degree to which the eudicot-based ABC model of flower organ identity applies to the other major subclass of angrosperms, the monocots, has yet to be fully explored. We cloned silky1 (si1), a male sterile mutant of Zea mays that has homeotic conversions of stamens into carpels and lodicules into palea/lemma-like structures. Our studies indicate that si1 is a monocot B function MADS box gene. Moreover, the si1 zag1 double mutant produces a striking spikelet phenotype where normal glumes enclose reiterated palea/lemma-like organs. These studies indicate that B function gene activity is conserved among monocots as well as eudicots. In addition, they provide compelling developmental evidence for recognizing lodicules as modified petals and, possibly, palea and lemma as modified sepals.     

花对称性的研究进展

花对称性(floral symmetry)是被子植物花部结构的典型特性之一,主要有辐射对称和两侧对称两种形式。被子植物初始起源的花为辐射对称,而两侧对称的花则是由辐射对称的花演变而来。两侧对称的花部结构是被子植物进化过程中的一个关键的革新,被认为是物种形成和分化的关键推动力之一。近年来有关花对称性的形成和进化机制的研究在植物学科的不同领域均取得了长足的进展。本文综述了花对称性在发育生物学、传粉生物学、生殖生态学及分子生物学等方面的研究进展。两侧对称形成于被子植物花器官发育的起始阶段,随后贯穿整个花器官发育过程或者出现在花器官发育后期的不同阶段。花器官发育过程中一种或多种类型器官的败育以及特异性花器官结构的形成是两侧对称形成的主要原因。研究表明,在传粉过程的不同阶段,花对称性均会受到传粉昆虫介导的选择作用。相比辐射对称的花,两侧对称的花提高了特异性传粉者的选择作用,增加了花粉落置的精确性,进而确保了其生殖成功。花对称性的分子机理已经在多种双子叶植物中进行了深入的研究。现有的证据表明,CYC同源基因在花对称性的分子调控方面起着非常重要的作用。花对称性在被子植物进化过程中是如何起源,与其他花部构成之间是否协同作用,一些不符合一般模式的科属其花对称性的形成机制等都是今后要进一步研究的命题。     

BEAK LIKE SPIKELET1 is Required for Lateral Development of Lemma and Palea in Rice

Lemma and palea are unique floral structures found only in Poaceae, and are responsible for protecting the inner floral organs and kernels from environmental stresses. However, the mechanism underlying specification of their morphology remains unclear. In this study, we characterized a rice mutant, beak like spikelet1 (bls1), which specifically affects development of the lemma and palea. In bls1 mutant, floral-organ identity and floral-organ patterning are normal, and the defects occur at the stage of the lemma and palea expansion, whereas the other aspects of floral architecture and form are not affected. We isolated BLS1 by positional cloning and found that it encodes a protein with a conserved domain of unknown function. BLS1 is expressed strongly in young inflorescence, specifically the young lemmas and paleas of spikelets. Subcellular localization analysis showed that BLS1 is localized in the nucleus. Expression of the AP1-like and SEP-like floral homeotic genes were not changed in the bls1 mutant. Our study suggested that BLS1 is required for lateral development of the lemma and palea and does not function at stages of floral-organ initiation and patterning.     

Molecular mapping of split rice spikelet mutant srs-1 and analysis of its homeotic function in rice

srs-1, a new floral organ identity gene in rice, was mapped using RAPD and RFLP markers. Firstly, the cross was made between "ZhaiYeQing 8" (ZYQ8, indica) and split rice spikelet (SRS, japonica) mutant. The ratio of wild-type individuals and mutant plants in F2 population is 3:1, which indicates that the mutant characteristics are controlled by single recessive gene, srs-1. Consequently, BSA method was adopted and 520 random 10-mer primers were used to screen polymorphic bands between two bulks. Six primers could amplify polymorphic bands, of which S465 generates the most stable RAPD patterns. Then, S465 that cosegregates in F2 population has been converted into an RFLP marker successfully. Furthermore, srs-1 gene was mapped on chromosome 3 using DH mapping population. The effect of srs-1 gene results in the mutant of split rice spikelet. The mutant has longer and softer palea/lemma than those of wild-type, and two small palea/lemma-like organs between palea and lemma. In addition, there is a flower with three stamens and carpel in the axil of lemma. Thus, there are nine stamens and two carpels in the spikelet of mutant. srs-1 gene may belong to homeotic gene of class A according to the mutant characteristics and ABC model.     

Molecular mapping of split rice spikelet mutantsrs-1 and analysis of its homeotic function in rice

srs-1, a new floral organ identity gene in rice, was mapped using RAPD and RFLP markers. Firstly, the cross was made between "ZhaiYeQing 8" (ZYQ8, indica) and split rice spikelet (SRS, japonica) mutant. The ratio of wild-type individuals and mutant plants in F2 population is 3:1, which indicates that the mutant characteristics are controlled by single recessive gene, srs-1. Consequently, BSA method was adopted and 520 random 10-mer primers were used to screen polymorphic bands between two bulks. Six primers could amplify polymorphic bands, of which S465 generates the most stable RAPD patterns. Then, S465 that cosegregates in F2 population has been converted into an RFLP marker successfully. Furthermore, srs-1 gene was mapped on chromosome 3 using DH mapping population. The effect of srs-1 gene results in the mutant of split rice spikelet. The mutant has longer and softer palea/lemma than those of wild-type, and two small palea/lemma-like organs between palea and lemma. In addition, there is a flower wit     

Identification and fine mapping of a mutant gene for palealess spikelet in rice

In grass, the evolutionary relationship between lemma and palea, and their relationship to the flower organs in dicots have been variously interpreted and wildely debated. In the present study, we carried out morphological and genetic analysis of a palealess mutant (pal) from rice (Oryza sativa L.), and fine mapping the gene responsible for the mutated trait. Together, our findings indicate that the palea is replaced by two leaf-like structures in the pal flowers, and this trait is controlled by one recessive gene, termed palealess1 (pal1). With a large F2 segregating population, the pal1 gene was finally mapped into a physical region of 35 kb. Our results also suggest that the lemma and palea of rice are not homologous organs, palea is likely evolutionarily equivalent to the eudicot sepal, and the pal1 should be an A function gene for rice floral organ identity.