Study of the inheritance of sex expression in the cucumber. The interaction of major genes with modifying genetic and non-genetic factors

Summary Two major genes and a complex of polygenes affecting sex expression inCucumis sativus L. as well as their interaction with some nongenetic factors were investigated in the present study. The genetic factorst was found to affect sex by inducing a shift of the predetermined flowering pattern of this plant, in the direction of its base. As this pattern is composed of a staminate stage followed by a mixed (staminate-pistillate) stage and a pistillate stage, a double dose ofst will induce a change from the normal monoecious sex expression (to absolute gynoecism.The second major gene studied,m, known previously to control sex in the individual flower (m/m—andromonoecious,M—monoecious), also interacts with factors affecting the flowering pattern by inducing male tendency.Using an ordinary commercial monoecious stock additional modifying factors for sex expression were demonstrated, by eight generations of selection for high (in male direction) and for low (in female direction) node number to the first pistillate flower. A comparison of the means and frequency distributions of node number of the two selected lines, their F₁ and F₂, indicated polygenic control of this character.It was found that two non-genetic factors, day length and gibberellic acid (GA), may mimic the genetic factors for sex expression. Furthermore, evidence was presented, indicating that this sex controlling ability of GA and the modifying genes may be based on physiological conditions common to those two genetic and nongenetic factors.The overall mechanism of sex control in the cucumber and in other plants was discussed and a hypothetic model for the evolution of dioecism in flowering plants was proposed.     

Preformation and distribution of staminate and pistillate flowers in growth units of Nothofagus alpina and N. obliqua (Nothofagaceae)

Background and Aims

The distribution and differentiation times of flowers in monoecious wind-pollinated plants are fundamental for the understanding of their mating patterns and evolution. Two closely related South American Nothofagus species were compared with regard to the differentiation times and positions of staminate and pistillate flowers along their parent growth units (GUs) by quantitative means.

Methods

Two samples of GUs that had extended in the 2004–2005 growing season were taken in 2005 and 2006 from trees in the Lanín National Park, Patagonia, Argentina. For the first sample, axillary buds of the parent GUs were dissected and the leaf, bud and flower primordia of these buds were identified. The second sample included all branches derived from the parent GUs in the 2005–2006 growing season.

Key Results

Both species developed flowering GUs with staminate and/or pistillate flowers; GUs with both flower types were the most common. The position of staminate flowers along GUs was similar between species and close to the proximal end of the GUs. Pistillate flowers were developed more distally along the GUs in N. alpina than in N. obliqua. In N. alpina, the nodes bearing staminate and pistillate flowers were separated by one to several nodes with axillary buds, something not observed in N. obliqua. Markovian models supported this between-species difference. Flowering GUs, including all of their leaves and flowers were entirely preformed in the winter buds.

Conclusions

Staminate and pistillate flowers of N. alpina and N. obliqua are differentiated at precise locations on GUs in the growing season preceding that of their antheses. The differences between N. alpina and N. obliqua (and other South American Nothofagus species) regarding flower distribution might relate to the time of anthesis of each flower type and, in turn, to the probabilities of self-pollination at the GU level.Key words: Branch, bud, growth unit, Markovian models, Nothofagus alpina, N. obliqua, Patagonian forests, pistillate flower, preformation, staminate flower     

Pollen movement to flowering canopies of pistillate individuals of three rain forest tree species in tropical Australia

This study measured the quantities of effective pollen vectors and their pollen loads arriving at the canopies of dioecious tropical rain forest trees in north-east Queensland. Population flowering synchrony, effective pollinator populations and pollen loads transferred between staminate and pistillate trees were compared among three insect-pollinated tree species. All three were visited by a wide range of insects, 75% of which (mostly 3–6 mm long) carried conspecific pollen. Fewer than 8% of individual insects were found to be carrying single-species pollen exclusively and none could be described as specialist pollen foragers. The introduced honeybee carried greater quantities of pollen than any native species but was not necessarily a reliable pollinator. The brief flowering periods in Neolitsea dealbata (3–4 weeks) and Litsea leefeana (4–5 weeks) populations were synchronized among individuals. Flowering in the Diospyros pentamera population extended over 15 weeks and most individuals were in flower for most of this period. Staminate trees began flowering earlier, produced more flowers and attracted relatively more insects than did pistillate trees, suggesting a density-dependent response of pollinators to flowering performance. Pollen was trapped in greater quantities on insects at staminate trees than at pistillate trees. Insect numbers increased at peak flowering periods and Diptera were the most abundant flower visitors. Anthophilous Coleoptera were more numerous at staminate than at pistillate trees in all three tree species populations. Larger quantities of pollen were mobilized during peak flowering times although the greatest quantities were transferred to pistillate canopies towards the end of the population flowering periods. Diptera carried pollen more often to pistillate N. dealbata and L. leefeana trees than did other groups whereas Coleoptera carried pollen more often to pistillate D. pentamera trees. The two contrasting flowering performances in the three tree species are discussed with reference to mechanisms that facilitate pollen transfer between staminate and pistillate trees.     

麻疯树雌雄花中MADS-BOX基因的表达分析

麻疯树(Jatropha curcas)种子含油率高,种子中的油类物质可作为生物柴油被开发和利用,是极具潜力的生物质能源树种之一。麻疯树雌雄异花,在自然条件下雄花数量通常远远大于雌花,这大大限制了种子和油的产量,因此开展麻疯树性别分化与花发育分子机理的研究具有重要意义。该研究选取10个麻疯树的MADS-BOX基因(JcAGL1,JcAGL6,JcAGL9,JcAGL11,JcAGL15,JcAGL61-3,JcAGL62-1,JcAGL62-6,JcAGL62-7,JcAGL80-2),提取麻疯树早期发育各个阶段的雌雄花总RNA,并反转录成cDNA,采用实时荧光定量方法,探索早期发育不同阶段的麻疯树雌雄花目的基因的表达情况。结果表明:目的基因在发育起始的雌雄花中的表达具有差异,比如JcAGL6和JcAGL15在雄花中表达量要高于雌花,而JcAGL1,JcAGL9和JcAGL11在雌花中的表达量要高于雄花,这说明花原基中目的基因表达会直接或间接决定性别分化的方向;在之后的发育过程中,目的基因的表达情况在雌雄花中有所不同:随着花的发育,目的基因在雌雄花中的表达量变化存在差别,这反应出麻疯树雌雄花发育中目的基因表达模式上的差异;另外,也能看出在此过程中各个目的基因又发挥着不同的功能。该研究结果为进一步探究麻疯树雌雄花发育相关基因的表达提供了理论依据,为了解麻疯树性别分化和花发育的分子机理奠定了基础。     

Flower structure and potential bisexuality in Freycinetia reineckei (Pandanaceae), a species of the Samoa Islands

In Freycinetia reineckei the staminate flower (on the staminate spikes) comprises 3 or 4 (sometimes 2) stamens and a pistillode with 2 (sometimes 4) carpellodes, and the pistillate flower (on the pistillate spikes) is formed of a pistil with 2 (sometimes 4) carpels and of 3 or 4 (sometimes 2) staminodes. This perfect floral homology, also observed in all the other species that were studied with both pistillate and staminate material, strongly suggests that the flower of Freycinetia is basically and potentially bisexual, and may explain the occasional sexual lability and bisexuality of that flower (occurrence of both pistillate and staminate inflorescences, and/or of bisexual inflorescences with bisexual flowers and/or unisexual flowers, on the same individuals) in some species, and also the frequent occurrence of bisexual spikes in this species. These may be partitioned into pistillate, staminate, mixed and sterile zones. In the pistillate zones the flowers have the same aspect and structure as the pistillate flowers. In the staminate zones the flowers generally comprise 3 or 4 (sometimes 2) stamens and a ‘semi-pistil’ some have both stamens and staminodes. The semi-pistils are intermediate between pistils and pistillodes in length, aspect and structure, but always have placentas and ovules. In the mixed zones the flowers are generally formed of a pistil and 3 or 4 (sometimes 2) stamens, and are therefore true hermaphrodite flowers; some have both stamens and staminodes. In the sterile zones the flowers comprise a semi-pistil and 3 or 4 (sometimes 2) staminodes. The staminodes are anatomically very similar to the stamens, especially in the staminate, mixed, and sterile zones, in which they exhibit a wide range of variation in length, aspect and structure. The perfect floral homology as generic character on one hand, and the occasional bisexuality both with and without bisexual flowers and other aspects of sex expression (e.g. occurrence of both pistillate and staminate shoots on the same individuals) in some species on the other hand, seem to indicate that Freycinetia is a basically monoecious, sex changing genus.     

Induction of male flowers in a pistillate line ofRicinus communis L. by silver and cobalt ions

Aqueous solutions of silver nitrate (10–100 g/plant) and cobalt chloride (125–500 g/plant), injected into the main stem of plants of the pistillate cv. 240 ofRicinus communis when the vegetative shoot apex was beginning to become reproductive, induced the formation of staminate (male) flowers with viable pollen in the normally strictly pistillate (female) terminal inflorescence, their number increasing with the dose of Ag⁺ and Co²⁺. No formation of bisexual flowers was noted. Female flowers pollinated with pollen from the induced male ones produced fruits and viable seeds.     

Resource partitioning to male and female flowers of Spinacia oleracea L. in relation to whole-plant monocarpic senescence

Male plants of spinach (Spinacea oleracea L.) senesce following flowering. It has been suggested that nutrient drain by male flowers is insufficient to trigger senescence. The partitioning of radiolabelled photosynthate between vegetative and reproductive tissue was compared in male (staminate) versus female (pistillate) plants. After the start of flowering staminate plants senesce 3 weeks earlier than pistillate plants. Soon after the start of flowering, staminate plants allocated several times as much photosynthate to flowering structures as did pistillate plants. The buds of staminate flowers with developing pollen had the greatest draw of photosynthate. When the staminate plants begin to show senescence 68% of fixed C was allocated to the staminate reproductive structures. In the pistillate plants, export to the developing fruits and young flowers remained near 10% until mid-reproductive development, when it increased to 40%, declining to 27% as the plants started to senesce. These differences were also present on a sink-mass corrected basis. Flowers on staminate spinach plants develop faster than pistillate flowers and have a greater draw of photosynthate than do pistillate flowers and fruits, although for a shorter period. Pistillate plants also produce more leaf area within the inflorescence to sustain the developing fruits. The (14)C in the staminate flowers declined due to respiration, especially during pollen maturation; no such loss occurred in pistillate reproductive structures. The partitioning to the reproductive structures correlates with the greater production of floral versus vegetative tissue in staminate plants and their more rapid senescence. As at senescence the leaves still had adequate carbohydrate, the resources are clearly phloem-transported compounds other than carbohydrates. The extent of the resource redistribution to reproductive structures and away from the development of new vegetative sinks, starting very early in the reproductive phase, is sufficient to account for the triggering of senescence in the rest of the plant.     

Experimental defoliation affects male but not female reproductive performance of the tropical monoecious plant Croton suberosus (Euphorbiaceae)

Background and Aims

Monoecious plants have the capacity to allocate resources separately to male and female functions more easily than hermaphrodites. This can be advantageous against environmental stresses such as leaf herbivory. However, studies showing effects of herbivory on male and female functions and on the interaction with the plant''s pollinators are limited, particularly in tropical plants. Here, the effects of experimental defoliation were examined in the monoecious shrub Croton suberosus (Euphorbiaceae), a wasp-pollinated species from a Mexican tropical dry forest.

Methods

Three defoliation treatments were applied: 0 % (control), 25 % (low) or 75 % (high) of plant leaf area removed. Vegetative (production of new leaves) and reproductive (pistillate and staminate flower production, pollen viability, nectar production, fruit set, and seed set) performance variables, and the abundance and activity of floral visitors were examined.

Key Results

Defoliated plants overcompensated for tissue loss by producing more new leaves than control plants. Production of staminate flowers gradually decreased with increasing defoliation and the floral sex ratio (staminate : pistillate flowers) was drastically reduced in high-defoliation plants. In contrast, female reproductive performance (pistillate flower production, fruit set and seed set) and pollinator visitation and abundance were not impacted by defoliation.

Conclusions

The asymmetrical effects of defoliation on male and female traits of C. suberosus may be due to the temporal and spatial flexibility in the allocation of resources deployed by monoecious plants. We posit that this helps to maintain the plant''s pollination success in the face of leaf herbivory stress.     

Honey bees as carriers of lychee erinose miteEriophyes litchii (Acari: Eriophyiidae)

Up to 23% of honey bees (Apis mellifera Linnaeus) collected from flowering lychee trees, severely infested with the lychee erinose mite (Eriophyes litchii Keifer), were found to be carrying live mites which were picked up as the bees foraged. This method of lychee erinose mite dispersal is discussed in the context of periodic records of spontaneous infestation of lychee flower panicles from previously uninfested orchards.     

核桃揪雌雄异熟性的初步研究

对核桃揪开花特性进行了观察。结果表明：核桃揪多数植株为雌雄异熟,有的植株上雌花先开（雌先型）,有的植株上雄花先开（雄先型）。每一植株上雌雄异熟的次序是相当稳定的。并且多数能为雌先型植株提供授粉机会的雄先型群体,其雌花期与雌先型群体的雄花期也能较好的吻合。但是少数花期不同的同一类型（即：雌先型与雌先型或雄先型与雄先型）植株之间也有授粉机会。在坐果率、产量、核果的整齐度、放叶早晚、落叶期、形态差异的调查分析中未发现异熟性与这些性状上有必然的联系。     

“Intra-reproductive vernalization” inSoldanella minima

Summary 1. The development of flower buds inSoldanella minima grown at 15° or above is stopped at the stage of sporogenous cells in the anthers. 2. Low temperature overcomes this inhibition, resulting in microsporogenesis and subsequent flowering. Microsporogenesis must thus be considered as a phase initiated only at low temperature. Photoperiod does not seem to playa decisive role in this process. 3. Applied gibberellic acid substitutes for the cold requirement of the flower buds.Dedicated to ProfessorR. Harder on the occasion of his 80. birthday.Research carried out under the auspices of the Consiglio Nazionale delle Ricerche (National Research Council, Italy).     

The control by gibberellic acid of stem elongation and flowering in biennial plants of Centaurium minus Moench

Summary Application of a small amount of gibberellic acid (GA₃) to unvernalized rosettes of a biennial strain of Centaurium minus Moench brings about immediate stem elongation under both long days (LD) and short days (SD), but the rate of stem elongation falls after the cessation of treatment. Under LD, but not SD, a second period of rapid and prolonged stem elongation may subsequently take place, associated with flowering. Extended GA₃ treatment under SD leads to the formation of a long stem but not to flowering; after the treatment the plants revert to vegetative aerial rosettes unless transferred to LD prior to the cessation of stem elongation; after such a transfer, rapid stem elongation and flowering may be initiated. If flower primordia are initiated under LD, stem elongation and formation of flower primordia continue after transfer to SD, though flowers do not develop fully. It is suggested that under LD but not SD applied GA₃ may bring about the production of endogenous gibberellin, and that this synthesis of endogenous gibberellin occurs in the flower primordia.     

Mineral nutrition and leaf longevity in an evergreen shrub,Ledum palustre ssp. decumbens

Summary Effects of variable mineral nutrient status on evergreen leaf longevity were investigated in a field fertilization experiment, and by comparison of plants from several sites. The species studied was Ledum palustre spp. decumbens, with a normal leaf life expectancy of 2.06 years and a maximum leaf longevity of 4.5–5.0 years. Most leaf losses took place during the growing season, not during the winter. Fertilization increased leaf production but decreased leaf survivorship. Total number of leaves per stem was unchanged with fertilization. In a comparison among sites, there was a moderate negative correlation between plant N and P concentrations and leaf longevity. These intraspecific responses are similar to known interspecific changes along nutrient gradients, i.e. with high nutrient availability a vegetation should become more deciduous and less evergreen.     

POLLINATION ECOLOGY OF A SEAGRASS,THALASSIA TESTUDINUM (HYDROCHARITACEAE), IN ST. CROIX

The reproductive ecology of a dioecious Caribbean seagrass, Thalassia testudinum was studied in St. Croix, U.S. Virgin Islands. Flowering in Thalassia coincided with spring tides with anthesis and anther dehiscence in staminate plants occurring at night. As predicted by search theory, pollen is dispersed in negatively buoyant rafts of pollen grains (that superficially resemble strings of frog spawn) which are bound by a slime of apparent thecal origin. Dispersal of pollen is submarine and approximately two-dimensional along the plane of the substratum. The stigmas on pistillate plants are linear, stiff, and densely papillate; they protrude from the mouth of the subtending leaf. Floral ratios of staminate to pistillate flowers were 60 to 1 with an average distance of 4.0 cm from pistillate flower to the closest staminate flowers. Populations included a high proportion of short shoots with numerous (> 100) leaf scars indicating ages for these axes of up to 3–4 years. Nonflowering shoots can be sexed and frequency of flowering can be assessed from scars of old inflorescences. Preliminary results suggest that there is no regular pattern of flowering, since the interval between flowering events is variable.     

单性木兰（木兰科）花的形态发生

报道了单性木兰(Kmeria septentrionalis Dandy)花的形态发生过程。发现过去一直被认为是雌花条状披针形的“内轮花被片”,实际为退化雄蕊,它形态发生的时间与位置均与雄花的雄蕊相同,在成熟结构中仍可见药室残迹,说明单性木兰的雌性花是由两性花退化而来。通过与K. duperreana(Pierre) Dandy和Magnolia thailandica Noot. &; Chalermglin雌花的比较,发现它们雌花的形态相同,从而得知人们长期以来对此3种植物雌花的认识有误,原一直认为的“内轮花被片”实为退化雄蕊。     

Subdioecy in Consolea spinosissima (Cactaceae): breeding system and embryological studies

The breeding system and the embryology of Consolea spinosissima, a tree-like opuntioid endemic to Jamaica, were investigated. Morphological and embryological studies revealed that the species is subdioecious, with three sexual morphs present in the 150 × 120 m plot studied at Hellshire Hills, Jamaica. The female morph has pistillate flowers with open stigma lobes, no pollen grains, and sets fruit. The male morph has cryptic staminate flowers with closed stigma lobes, viable pollen grains, and a nonfunctional gynoecium that does not set seed. The weak hermaphrodite morph has low fruit set and "perfect" flowers that superficially resemble the functionally staminate flowers of the male morph. These perfect flowers reach anthesis with viable pollen grains, with no or only a few functional ovules, and with the style supporting pollen tube growth. Embryological studies showed that the critical stage for sex determination occurs earlier in pistillate than in staminate and perfect flowers. Anthers of pistillate flowers abort prior to microspore tetrad formation, whereas ovules of the staminate and perfect flowers degenerate after the complete maturation of the embryo sac. Based on flower structure and embryological data, we hypothesize that the ancestor of C. spinosissima is/was hermaphroditic.     

Effects of L-Glutamate Transport Inhibition by a Conformationally Restricted Glutamate Analogue (2S,1'S,2'R)-2-(Carboxycyclopropyl)Glycine (L-CCG III) on Metabolism in Brain Tissue In Vitro Analysed by NMR Spectroscopy

(2S,1'S,2'R)-2-(Carboxycyclopropyl)glycine (L-CCG III) was a substrate of Na⁺-dependent glutamate transporters (GluT) in Xenopus laevis oocytes (IC₅₀ 13 and 2 M for, respectively, EAAT 1 and EAAT 2) and caused an apparent inhibition of [³H]L-glutamate uptake in mini-slices of guinea pig cerebral cortex (IC₅₀ 12 M). In slices (350 M) of guinea pig cerebral cortex, 5 M L-CCG III increased both the flux of label through pyruvate carboxylase and the fractional enrichment of glutamate, GABA, glutamine and lactate, but had no effect on total metabolite pool sizes. At 50 M L-CCG III decreased incorporation of ¹³C from [3-¹³C]-pyruvate into glutamate C4, glutamine C4, lactate C3 and alanine C3. The total metabolite pool sizes were also decreased with no change in the fractional enrichment. Furthermore, L-CCG III was accumulated in the tissue, probably via GluT. At lower concentration, L-CCG III would compete with L-glutamate for GluT and the changes probably reflect a compensation for the missing L-glutamate. At 50 M, intracellular L-CCG III could reach > 10 mM and metabolism might be affected directly.     

1H, 15N and 13C resonance assignments and monomeric structure of the amino-terminal extracellular domain of epithelial cadherin

Summary E-cadherin is a transmembrane protein that provides Ca²⁺-dependent cell adhesion to epithelial cells. The large majority of the ¹H, ¹⁵N, ¹³C and ¹³CO resonances of a 146-amino acid polypeptide from epithelial (E-) cadherin have been assigned using multidimensional NMR spectroscopy. The structure of the amino-terminal 100 amino acids, corresponding to the first extracellular repeat of E-cadherin [Overduin et al. (1995) Science, 267, 386–389], has been refined. The monomeric state of this isolated domain is demonstrated by light scattering and sedimentation analysis. Seven -strands and two short helices were identified by patterns of NOE cross-peaks, vicinal coupling constants and chemical shift indices. A novel structural motif termed a quasi--helix found in the crystal structure of a neural (N-) cadherin domain [Shapiro et al. (1995) Nature, 374, 327–337] is characterized in detail for the first time by NMR. Slowly exchanging amides were concentrated in the -sheet region and quasi--helix. The -barrel fold of the cadherin domain is topologically similar to the immunoglobulin fold. Comparison of this solution structure to the crystallized dimers of the N-terminal pair of E-cadherin domains [Nagar et al. (1996) Nature, 380, 360–364] and of the homologous single domain of N-cadherin reveals a conserved cadherin fold with minor structural differences, which can be accounted for by differences in metal ligation and oligomeric state.Abbreviations cad extracellular cadherin repeat - CAM cell adhesion molecule - CSI chemical shift index - DTT dithiothreitol - E-cadherin epithelial cadherin - N-cadherin neural cadherin - NOE nuclear Overhauser enhancement - PFG pulsed field gradient - rmsd root-mean-square deviation     

Gene expression in Mercurialis annun flowers: In vitro translation and sex genotype specificity. Male-specific cDNA cloning and hormonal dependence of a corresponding specific RNA

Summary Sequences specifically expressed in flowers of each sex type of Mercurialis annua have been demonstrated by a comparative analysis of the translation products of their poly(A)⁺ RNA populations (wheat germ system, two-dimensional electrophoresis). This method confirms previous results of hybridization kinetics: the staminate flowers of the normal fertile male (wild type) and restored fertile male strain (identical morphology) and those of the sterile male genotype contain specific poly(A)⁺ RNAs and sequences shared by sets of two of these males, as well as numerous common sequences. The pistillate flowers of the constructed female strain 19_–5 (carrying male sterility determinants) also contains specific poly(A)⁺ RNA compared to identical flowers of a normal female genotype. In vitro translation, however, showed a specificity (not revealed by hybridization kinetics) in the normal female genotype compared to a normal male genotype and to the female strain 19_–2 (female strain cDNAs hybridized 100% to poly(A)⁺ RNAs of male and female 19_–5 genotypes). Cloning certain specific sequences (cDNAs) of the fertile male wild type and using one cDNA as probe also confirms the previously described male specificity. Moreover, the hormonal dependence of RNA corresponding to a specific male probe is demonstrated: its kinetics of disappearance are a function of the action of the feminizing hormone (cytokinins). These results are in agreement with our hypothesis of sexual organogenesis: sexual hormones controlled by regulator genes of the sexual genotype induce, in definite cell lineages of bipotential meristems, the expression of genes specific for male or female expression.