Floral development and the formation of unisexual spikelets in the Andropogoneae (Poaceae)

We investigated spikelet development in four distantly related species of the grass tribe Andropogoneae to determine whether spikelet development and the formation of unisexual florets are uniform throughout the tribe. We studied development in Bothriochloa bladhii, Coelorachis aurita, Heteropogon contortus, and Hyparrhenia hirta, and compared these with Panicum, a member of the sister tribe Paniceae. Many aspects of spikelet development in the species we have studied correlate with what is already known for Tripsacum and maize (both Andropogoneae), despite variation in how unisexual florets are distributed on the plant. The formation of unisexual spikelets is also uniform. All florets initiate both pistil and stamen primordia. In florets destined to be male, cell death occurs in the subepidermal layers of the gynoecium after the formation of a gynoecial ridge. In florets destined to be female, there is no apparent cell death in the stamens, but growth ceases after anther formation. The similarity in spikelet development and the formation of unisexual florets point to a common genetic mechanism for sex determination throughout the Andropogoneae and possibly the entire Panicoideae. Use of a cell death pathway to cause gynoecial abortion may be the basis of one morphological character that defines the subfamily.     

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.     

Development of Male Flowers in Zizania aquatica (North American Wild-Rice; Gramineae)

We investigated the histology and developmental morphology of flowers of wild-rice (Zizania aquatica), a member of grass subfamily Oryzoideae, to determine whether male flowers in this species develop in a manner similar to those in the subfamily Panicoideae, a group that includes many species with unisexual flowers. Zizania has evolved unisexual flowers from hermaphrodite ancestors and is only distantly related to the Panicoideae; the origins of unisexual flowers are independent in the two taxa. Ontogenetic evidence indicates that many species within the subfamily Panicoideae develop male flowers by a process similar to that already described for maize (Zea mays), a panicoid grass. Unisexual male flowers in maize initiate both the stamen (androecium) and the pistil (gynoecium), but the gynoecium aborts early in development through the death of the subepidermal cells. Cell death in gynoecia of maize is known to be controlled by the product of the gene tasselseed2 (ts2), and an orthologue of ts2 has been shown to have the same effect in the sister genus Tripsacum. It seems likely that ts2 orthologues mediate cell death throughout the Panicoideae, but the phylogenetic range of the cell death mechanism is not known. In this study we show that male flowers of Z. aquatica show neither the distinctive pattern of cell death nor the ontogenetic timing of abortion that are characteristic of male flower formation in studied species of Panicoideae. This indicates that these unisexual flowers may be produced by an entirely different mechanism from that employed by the Panicoideae. Either ts2 does not control sex expression in Zizania, or it is deployed at a different time, and possibly in different tissues, with a different histological result. Our results indicate that the independent origins of male flowers in Gramineae apparently do not have a common system of genetic control.     

Cell cycle arrest of stamen initials in maize sex determination

The maize sex determination pathway results in the arrest of stamen in ear spikelets and the abortion of pistils in both the tassel spikelets and in the secondary florets of ear spikelets. Arrested stamen cells showed no signs of DNA fragmentation, an absence of CYCLIN B expression, and an accumulation of the negative cell cycle regulator WEE1 RNA.     

Senescence-induced expression of cytokinin reverses pistil abortion during maize flower development

Maize is a monoecious species that produces imperfect (unisexual), highly derived flowers called florets. Within the spikelet, the basic repeating unit of the maize inflorescence, the spikelet meristem gives rise to an upper and a lower floret. Although initially bisexual, floret unisexuality is established through selective organ elimination. In addition, the lower floret of each ear spikelet is aborted early in its development, leaving the upper floret to mature as the only pistillate floret. Expression from the cytokinin-synthesizing isopentenyl transferase (IPT) enzyme under the control of the Arabidopsis senescence-inducible promoter SAG (senescence associated gene)12 was observed during early maize floret development. Moreover, the lower floret was rescued from abortion, resulting in two functional florets per spikelet. The pistil in each floret was fertile, but the spikelet produced just one kernel composed of a fused endosperm with two viable embryos. The two embryos were genetically distinct, indicating that they had arisen from independent fertilization events. These results suggest that cytokinin can determine pistil cell fate during maize floret development.     

Systematics and Evolution of Subfam. Panicoideae (Gramineae)

Subfam. Panicoideae consists of 230 genera and 3000 species, about 1/3 of the total number of Gramineae. The evolution in Gramineae (Poaceae) is considered to have been proceeding in the direction of simplification. Three major lineages in Panicoideae are recognized based on analysis of characters, geographical distribution and ecological preferences. They are represented respectively by three supertribes i.e. Panicatae, Andropogonatae and Maydatae. The first supertribe has well-developed florets, fertile lemma indurate, but is awnless and thus is the primitive group in the subfamily. In the second supertribe spikelets are well-developed, the lower glume is larger than the florets and embraces them, the florets are awned. Particularly significant is the evolution of spikelets from solitary to paired, from the two spikelets of each pair both bisexual and alike in form to heterogamous pair of spikelets in which the sessile one is fertile and the pedicelled one sterile and further to homogamous pair of spikelets at base of inflorescence. In this lineage occurs an inflorescence protected by a large sheathing bract. It is the most flourishing by developed evolutionary branch. In the third supertribe raches are developed, thick, inflorescences have become pillar-shaped with spikelets sunken in the raches. The reduction of spikelets from bisexual to unisexual has taken place and they occur on separte inflorescenes, bringing about subsequent sexual differentiation of inflorescences. Raches have also evolved from solitary to united. The supertribe Maydatae is likely to be one of the most advanced group in Gramineae. Subfam. Panicoideae occurs in the tropics and subtropics, and these three supertribes have apparently followed three lines of development under different environmental conditions. The supertribe Panicatae adapts to hot moist conditions, the supertribe Andropogonatae adapts to a monsoon climate, occurring in savannas, the supertribe Maydatae occurs in monsoon moist conditions.     

菰属系统与演化研究——孕花外稃表皮微形态

应用扫描电子显微镜对禾本科菰属及其有关9属14种3变种1变型的孕花外稃顶端及其芒基部的表皮微形态进行观察比较研究。研究结果基本符合E.E.Terrell与W.P.Wergin的研究报道,但我们不仅种类收集齐全而且增加研究了与菰属有关的属种,及菰属下的一些变种及混淆种,更有利于探讨菰属的系统位置及其演化关系。菰属以其稻型的双胞微毛与硅质体应隶属于稻族。又根据它们所具硅质体形状及硅质乳突的数量,除揭示它为单性花一支与稻属、假稻属、水禾属等两性花一支在稻族中存在平行演化现象外,也揭示了在菰属中共有4种、2亚种,其中以菰最为原始。     

DEVELOPMENT OF TASSEL SEED 2 INFLORESCENCES IN MAIZE

The normal pattern of maize floral development of staminate florets on the terminal inflorescence (tassel) and pistillate florets on the lateral inflorescences (ears) is disrupted by the recessive mutation tassel seed 2. Tassel seed 2 mutant plants develop pistillate florets instead of staminate florets in the tassel. In addition, the ears of tassel seed 2 plants display irregular rowing of kernels due to the development of the normally suppressed lower floret of each spikelet. The morphology of tassel and ear florets of the recessive maize mutant tassel seed 2 has been compared to those of wild-type maize through development. We have identified the earliest stages at which morphological signs of sex differentiation are evident. We find that sex determination occurs during the same stage on tassel and ear development. Early postsex determination morphology of florets in wild-type ears and in tassel seed 2 tassels and ears is identical.     

Unisexual flower, spikelet, and inflorescence development in monoecious/dioecious Bouteloua dimorpha (Poaceae, Chloridoideae)

Unisexual flowers have evolved repeatedly in the angiosperms. In Poaceae, multiple transitions from bisexual to unisexual flowers are hypothesized. There appear to be at least three distinct developmental mechanisms for unisexual flower formation as found in members of three subfamilies (Ehrhartoideae, Panicoideae, Pharoideae). In this study, unisexual flower development is described for the first time in subfamily Chloridoideae, as exemplified by Bouteloua dimorpha. Scanning electron microscopy (SEM) and anatomy were used to characterize the development of male (staminate) and female (pistillate) flowers, spikelets, and inflorescences. We found the developmental pathway for staminate flowers in B. dimorpha to be distinct from that described in the other three subfamilies, showing gynoecial arrest occurs at a different stage with possible loss of some cellular contents. However, pistillate flowers of B. dimorpha had some similarity to those described in other unisexual-flowered grasses, with filament and anther differentiation in abortive stamens. Comparing our findings with previous reports, unisexual flowers seem to have evolved independently in the four examined grass subfamilies. This analysis suggests the action of different genetic mechanisms, which are consistent with previous observations that floral unisexuality is a homoplasious condition in angiosperms.     

Genetic variability is correlated with population size and reproduction in American wild-rice (Zizania palustris var. palustris, Poaceae) populations

American wild-rice (Zizania palustris var. palustris) has served as a staple for indigenous North Americans for thousands of years, but has had significant habitat losses in recent centuries. We investigated genetic variability among 17 wild-rice populations in northern Wisconsin using 13 isozyme markers. We then compared these genetic patterns to differences in habitat and population characteristics and phenotypic variation in plant growth and reproduction across sites. Wild-rice's mean genetic diversity (0.15) is moderate compared to wind-pollinated outcrossers but lower than the mean (0.20) reported for the Poaceae. Estimated inbreeding coefficients within populations (f) average 0.12 but vary greatly among the populations (from -0.44-0.52), suggesting heterogeneous population histories. Larger populations in larger lakes express higher levels of genetic variability and smaller inbreeding coefficients than smaller or more isolated populations. The number of panicles per plant is also higher in populations with greater genetic variability. Estimated genetic differentiation among the 17 populations (F(ST)) was high (0.30), suggesting limited gene flow among drainages. Wild-rice population size and degree of isolation have opposing effects on its genetic variability, and plant performance is positively associated with genetic variability.     

Sex Determination by Sex Chromosomes in Dioecious Plants

Abstract: Sex chromosomes have been reported in several dioecious plants. The most general system of sex determination with sex chromosomes is the XY system, in which males are the heterogametic sex and females are homogametic. Genetic systems in sex determination are divided into two classes including an X chromosome counting system and an active Y chromosome system. Dioecious plants have unisexual flowers, which have stamens or pistils. The development of unisexual flowers is caused by the suppression of opposite sex primordia. The expression of floral organ identity genes is different between male and female flower primordia. However, these floral organ identity genes show no evidence of sex chromosome linkage. The Y chromosome of Rumex acetosa contains Y chromosome-specific repetitive sequences, whereas the Y chromosome of Silene latifolia has not accumulated chromosome-specific repetitive sequences. The different degree of Y chromosome degeneration may reflect on evolutionary time since the origination of dioecy. The Y chromosome of S. latifolia functions in suppression of female development and initiation and completion of anther development. Analyses of mutants suggested that female suppressor and stamen promoter genes are localized on the Y chromosome. Recently, some sex chromosome-linked genes were isolated from flower buds of S. latifolia.     

RELATIONSHIP BETWEEN MALE AND FEMALE GAMETOPHYTE DEVELOPMENT IN RYE

The development of male gametophyte and female gametophyte within a floret of rye (Secale cereale L.) was examined. Generally, meiosis in microsporocytes and in megasporocytes occurs simultaneously in most florets, but the period from zygotene to tetrad meiosis in the megasporocyte progresses more slowly than that in the microsporocyte. When the female gametophyte has one nucleus and no vacuoles, the male gametophyte has a single, eccentric nucleus. By the time the female gametophyte develops to the vacuolated one-, two-, four-, and eight-nucleate stages and to the growth and differentiation of the egg apparatus stage, the male gametophyte reaches the two-celled pollen stage. As the female gametophyte matures, the male gametophyte also reaches maturity. The duration of male gametophyte development from microspore mother cell and the duration of female gametophyte development from megaspore mother cell are the same in most florets. The relationship between sexual development of cross-pollinated rye is similar to that of self-pollinated wheat (Triticum aestivum L.). It seems that the relationship is not related to the breeding system.     

Floret Initiation and Development in Spring Wheat (Triticum aestivum L.)

The number and developmental stages of florets were determinedin each spikelet of the spike in the main shoots of spring wheat.Samples were taken frequently from plants grown in a phytotronand in a nitrogen application field-test. Ten stages of development,from floret initiation until anthesis, were recognized and described. Inter-spikelet variation in the total number of initiated floretswas rather small. However, the number of florets at advancedstages of development, as well as the number of grains, washighest in the central spikelets in which florets initiatedfirst. Floret initiation did not proceed beyond spike emergence,whereafter the distal florets and the spikelet apex degenerated.Grain-set was restricted to florets which had developed at leastto the stage of visible anther lobes at spike emergence. Thenumber of these florets was increased significantly by nitrogenapplication. Wheat, Triticum aestivum L., spikelet, floret, grain set, nitrogen     

Function and evolution of sterile sex organs in cryptically dioecious Petasites tricholobus (Asteraceae)

Background and Aims

Why are sterile anthers and carpels retained in some flowering plants, given their likely costs? To address this question, a cryptically dioecious species, Petasites tricholobus, in which male and female plants each have two floret types that appear pistillate and hermaphroditic, was studied. The aim was to understand the function of sterile hermaphroditic florets in females. In addition, the first examination of functions of sterile female structures in male plants was conducted in the hermaphroditic florets on males of this species. These female structures are exceptionally large in this species despite being sterile.

Methods

Differences in floret morphology between the sex morphs were documented and the possible functions of sterile sex organs investigated using manipulative experiments. Tests were carried out to find out if sterile female structures in male florets attract pollinators and if they aid in pollen dispersal, also to find out if the presence and quantity of sterile hermaphroditic florets in females increase pollinator attraction and reproductive success. To investigate what floret types provide nectar, all types of florets were examined under a scanning electron microscope to search for nectaries.

Key Results

The sterile female structures in male florets did not increase pollinator visits but were essential to secondary pollen presentation, which significantly enhanced pollen dispersal. Sterile pistillate florets on male plants did not contribute to floral display and disappeared in nearly half of the male plants. The sterile hermaphroditic florets on female plants attracted pollinators by producing nectar and enhanced seed production.

Conclusions

The presence of female structures in male florets and hermaphroditic florets on female plants is adaptive despite being sterile, and may be evolutionarily stable. However, the pistillate florets on male plants appear non-adaptive and are presumably in decline. Differential fates of the sterile sex organs in the species are determined by both the historical constraints and the ecological functions.     

Effects of exogenous hormones on floret development and grain setin wheat

At specific stages during floret development, solutions of IAA,GA₃, zeatin and ABA were injected into the leaf sheath around theyoung spike of wheat (Triticum aestivum L.) to study theregulating effects of exogenous hormones on floret development. Zeatin promotedfloret development and significantly increased the number of fertile florets aswell as grain set, especially at the stage of anther-lobe formation. Zeatinalsoincreased the sugar concentrations in spikes at anthesis. In contrast, IAA,GA₃ and ABA inhibited floret development, with different patternsforeach of the hormones. IAA inhibited the development of the whole spike and allflorets in the spikelets such that grain loss occurred in all positions in thespikelets. GA₃ increased the number of fertile florets per spike,butdecreased grain set of the third floret in each spikelet, especially whenapplied at terminal spikelet formation. ABA inhibited floret development, anddecreased the number of fertile florets and grain set at almost all developmentstages, except at anther-lobe formation. The inhibitory effect of ABA wasmainlyon the first and third florets in each spikelet.     

Changes in Cell Number and Cell Division Activity during Endosperm Development in Allohexaploid Wheat, Triticum aestivum L.

Nuclear or cell number, and the mitotic index, were recordedin endosperms of Triticum aestivum cv. Mardler to test if aparticular stage of endosperm development was critical in determiningthe final grain weight. The basal four florets of emasculatedspikelets (controls), and the third and fourth florets of spikeletswhere the two basal ovaries were removed (ovary-removed), weresampled at various times up to 360 h after hand-pollination.The coenocytic phase in endosperms ended about 84 h after pollinationregardless of both grain position and the treatment. The onsetof the cellular stage was characterized by the final large fluctuationsin the mitotic index reflecting the culmination of the synchronousnuclear division of the coenocytic stage. Thereafter, the mitoticindex fluctuated with smaller amplitudes and, by 216 h afterpollination, was < 1%. Neither floret position in the spikeletnor the treatment affected the pattern of alteration to themitotic index. However, ovary removal from first and secondflorets resulted in significantly heavier grains and higherendosperm cell number in the 3rd and 4th florets compared withthe controls. In all florets, mean endosperm cell number peakedat 280 h but decreased by 360 h after pollination. At this time,the mean cell numbers in endosperms of the 3rd and 4th floretsof ovary-removed spikelets were significantly higher than inthe corresponding endosperms in the controls. Thus, a key contributoryfactor in determining the final endosperm cell number may bethe number of cells which are lost during the late period ofthe cellular stage of endosperm development. Key words: Endosperm cell number, florets, grain weight, mitotic index, Triticum aestivum     

A Study of Floret Development in Wheat (Triticum aestivum L.)

Plants of wheat (Triticum aestlvum L.) cv. Aotea were grownat high or low nitrogen levels and in a natural photoperiodor continuous light. Starting 17–21 days from the double-ridgestage, eight plants from each treatment were sampled every 3days until anthesis, and the two basal, the sixth, and the terminalspikelets were sectioned longitudinally. A developmental scorewas assigned to each floret and rates of development calculated.Continuous light hastened development but reduced the numberof spikelets per ear, while high nitrogen delayed developmentbut increased spikelet numbers. The number of florets initiatedin each spikelet varied within narrow limits, but grain settingdepended strongly on spikelet position and on treatment. Althoughflorets were initiated in acropetal succession, the rate ofdevelopment tended to increase up to floret 4 but then declinedmarkedly. As a result grain setting was confined to basal floretpositions, although the two basal spikelets developed so slowlythat they contributed relatively little to grain yield. Distalflorets degenerated almost simultaneously at or before ear emergence,but those in intermediate positions continued to develop untilafter fertilization in the lower florets. It is argued thatthe spikelet is an integrated system in which correlative mechanismsplay a part throughout the development of the florets.     

Characterization of the common wheat (Triticum aestivum L.) mutation line producing three pistils in a floret

In a normal wheat (Triticum ssp.L.) spike, one floret carries only one pistil that will further develop into one grain after fertilization. The cultivated common wheat (T. aestivum L.) mutation line Three Pistils (TP) carried three pistils in a floret. Although one or two of the pistils died out before seed set in some florets, there were exist many florets that set three seeds. Normally, it was observed that there were one to three seeds in different florets of the same spike. Therefore, this mutation trait could raise considerably the number of grains per spike. The weight of 100 grains in three seeds set florets was lower than that of in one seed set florets. But three seeds set florets were significantly to surpass the one seed set florets in grain(s) weight per floret. Based on these results, the three pistils trait was suggested to be an interesting germplasm resource. Localisation of the gene controlling the three pistils trait was carried out by the method of crossing TP with the Chinese Spring disomic substitutions. F2 population segregation analysis revealed that only the 5B F2 population did not show homogeneity to control population. chi2-test analysis indicated that 5B F2 population, and only this population, was deviated from the Mendelian segregation ratio (3:1). As a conclusion, the gene for three pistils trait was located on chromosome 5B. According to the Recommended rules for gene symbolization in wheat, the name of the dominant gene for three pistils trait in the line TP was suggested as Pis1.     

苔草属植物的返祖两性小穗

本文报告了大别苔(Carex dabiecnsis)和白鳞苔草(C.polyschoena)的返祖小穗。这些返祖小穗中的类型之一具有与篙草属(Kobresia)以及Schoenoxiphium属完全相似的两性结构小穗,在这一类型的小穗中,它们生有一雌花及其在上的雄花。如图所示,大别苔和白鳞苔草中第(3)类型的返祖小穗和Schoenoxiphium属以及嵩草属的第1、2类型的小穗基本上没有区别,这些返祖小穗的发现证明果胞是苔草属的真正小穗,而且可以证明现存苔草的单性小穗是起源于具两性小穗的祖先。因此,可以相信具有两性结构的返祖小穗的发现进一步揭示了苔草亚科中Schoenoxiphium属,蒿草属,钩轴莎属和苔草属之间的系统发育的亲缘关系。