The relationship between xylem differentiation and dormancy evolution in apricot flower buds (Prunus armeniaca L.): the influence of environmental conditions in two Mediterranean areas

Environmental and climatic conditions affect the flower bud growth, flowering and yield performance of fruit species. Temperature and water availability appear to be important factors for temperate fruit trees in mild climates. The aim of this research was to study the involvement of temperature regime on biological processes, such as xylem differentiation and dormancy evolution, in apricot flower buds (Prunus armeniaca L.). Over 3 consecutive years, biological and anatomical investigations were carried out in Tuscany (Italy) and Murcia (Spain) on two cultivars characterized by a different chilling requirement (CR): Currot, a traditional Spanish cultivar with a very low CR, and Stark Early Orange (SEO), a North American cultivar with a very high CR. Currot had a regular bearing, and was characterized by a synchronism between endodormancy release and xylem differentiation. On the other hand, SEO showed an irregular flower bud growth leading to flowering and fruit-set failure, even with a high level of chilling accumulation. No relationship was found in SEO between xylem development and flower bud growth reactivation. The de-synchronism between these processes could determine the poor adaptability to different environmental areas of SEO cultivar, and its very high CR is not the main hypothesised cause of the altered flower bud development.     

Flower bud differentiation and development in fruiting and non-fruiting shoots in relation to fruit set in apricot (Prunus armeniaca L.)

Situations of high flower bud drop and low fruit set without apparent causes are common in fruit trees. The term flower quality has been coined to explain differences among flowers in their capacity to set fruit, but the causes underpinning these differences are largely unknown. This lack of knowledge is based on the fact that these differences are established a posteriori and there are no criteria to determine a priori what will make a flower to set a fruit or to drop. In this work, we profit from the empirical knowledge that there are fruiting and non-fruiting shoots to explore to which extent flower bud differentiation and bud development will affect the subsequent fruit set. For this purpose, the processes from flower bud differentiation to fruit set were sequentially analyzed in both types of shoots, over 2 years. More than half of the buds from long shoots aborted development and dropped before flowering. At anthesis, most of the remaining flowers showed underdeveloped pistils that failed to sustain pollen germination or pollen tube growth along the pistil. This unsuccessful development resulted in clear differences in fruit set between both types of branches. These results highlight that flower bud differentiation and development play an important role for fruit set and that developmental timing appears critical to reach anthesis with a fully developed pistil.     

常春二乔玉兰春夏季开花节律及营养效应研究

为系统掌握常春二乔玉兰春夏季开花物候节律,探讨其与营养物质的关系,本研究以6年生常春二乔玉兰为试验材料,观测其年生长发育节律、春夏季开花物候特性以及茎段营养物质的含量变化。结果表明：（1）每年12月始至翌年2月下旬为常春二乔玉兰休眠期。2月下旬花芽膨大生长,并于3月开始春季开花,花期持续约20 d。4月进行营养生长,5月完成花芽分化。5月底部分花芽膨大并于6月开始开花,夏季花期持续约20 d。7~9月为未膨大花芽的发育滞缓期。此外,少量夏季开放的花的基部侧芽再次分化形成花芽。10~12月随着落叶的开始,树体逐渐进入休眠期。（2）常春二乔玉兰营养生长后分化的花芽能够花开两季。春季开花为先花后叶,开花率为100%,开花同步率较高,雌、雄蕊发育正常,为可育花。夏季开花为花叶同放,开花率约为30%,且开花同步率较低,开放的花内雌、雄蕊发育异常,为不育花。（3）春季开花期间可溶性糖和可溶性蛋白呈下降趋势,淀粉含量于开花后期下降;夏季开花期间可溶性糖和淀粉总体呈先降后升趋势,而可溶性蛋白总体呈下降趋势。综上所述,常春二乔玉兰春、夏季开花期内开花模式存在一定差异,其显著节律特征与营养物质含量变化有关,推测低水平的可溶性糖及高水平的淀粉和可溶性蛋白有利于春季开花的启动,而低水平的可溶性蛋白及高水平的可溶性糖和淀粉含量则有利于夏季开花的实现。     

藤本月季“安吉拉”花芽分化形态结构及内源激素变化研究

以藤本月季“安吉拉”为试验材料,通过石蜡切片、体视显微镜观察及内源激素的测定,研究花芽分化过程的形态结构及内源激素的变化,为花期调控、景观品质的提升及相关育种工作提供基础数据。藤本月季“安吉拉”花芽各部分分化顺序由外向内进行,分为5个时期,共历时30 d,首先是生长锥呈圆锥状突起的形态分化期;扁平生长锥周围出现5个突起,即为萼片原基分化期;萼片原基的内方分化出成轮状的多个花瓣原基,即为花瓣原基分化期;花瓣原基基部从上向下分化出多轮雄蕊原基,即为雄蕊原基分化期;扁平的生长锥顶端突起形成多个雌蕊原基,为雌蕊原基分化期。随花芽分化进程脱落酸(ABA)、细胞分裂素(CTK)浓度变化规律相似,均呈先升高再下降趋势,萼片原基分化期其浓度均显著高于其他时期;生长素(IAA)浓度呈逐渐上升的趋势;赤霉素(GA)浓度呈逐渐下降趋势。随花芽分化进程IAA/GA和IAA/ABA比值呈逐渐上升趋势,CTK/GA和(ABA+CTK)/GA比值在萼片原基分化期显著高于形态分化期。内源激素测定结果表明:ABA、CTK浓度在萼片原基分化期显著升高与花芽分化诱导有关,较低的IAA浓度以及GA浓度的降低有利于藤本月季“安吉拉”的花芽分化;萼片原基分化期CTK/GA和(ABA+CTK)/GA比值升高可能与花芽分化诱导有关,高水平的IAA/ABA和IAA/GA比值可能与花器官原基的进一步发育相关。     

‘圣诞快乐’朱顶红花芽分化研究

采用解剖观测和石蜡切片技术,对朱顶红品种‘圣诞快乐’花芽生长情况、花器官分化和性细胞分化过程进行了研究,以明确朱顶红花芽分化特征,为其花发育、花期调控、杂交育种和系统分类研究提供理论依据。结果表明：‘圣诞快乐’朱顶红每年产生2个花序芽,在第2年完成其内花芽花器官分化,经过低温作用后于第3年盛开,其中第2个花序偶有败育发生;花器官分化过程包括花原基分化期、外花被原基分化期、内花被原基分化期、雄蕊原基分化期、心皮原基分化期,对应的花芽大小分别约为0.02、0.05、0.1、0.2、0.3 cm,所有花器官均为螺旋状向心式发生;朱顶红花药4室,花药壁从外至内由表皮、药室内壁、中层和绒毡层组成,绒毡层类型为分泌型,小孢子减数分裂类型为连续型,四分体排列方式为十字交叉型,成熟花粉粒为2-细胞型;朱顶红雌蕊3心皮,下位子房,中轴胎座,3心室,每室两列倒生胚珠,胚珠为双珠被,厚珠心,具葱型胚囊。     

敦煌‘李光杏’花芽发育过程中雌蕊败育观察及生物学特性调查

为克服杏生产中雌蕊败育现象、提高坐果率,该研究以甘肃省敦煌市的‘李光杏’盛果期不同树势(强势树、中庸树和弱势树)和不同类型结果枝(花束状果枝、短果枝、中果枝、长果枝)的花芽为试材,对其开花物候期及生物学特性进行调查,采用石蜡切片观察易发生败育时期(花芽分化初期、雌蕊分化期、花粉细胞期和花蕾膨大期)的花芽内部组织结构形态,同时测定各主要败育时期叶片可溶性糖、淀粉含量及矿质元素的变化。结果表明:(1)弱树花芽的平均败育率最高(91.26%),中庸树最低(71.08%);不同类型结果枝中,花束状果枝花芽的败育率最低,长果枝的败育率最高。(2)不同结果枝类型的花粉生活力表现为花束状果枝>短果枝>中果枝>长果枝,花粉发芽率表现为短果枝>花束状果枝>中果枝>长果枝。各花型花粉生活力表现为雌蕊高于雄蕊>雌雄等长>雌蕊低于雄蕊>无雌蕊,发芽率的表现为雌雄等长>雌蕊高于雄蕊>雌蕊低于雄蕊>无雌蕊。(3)各树势花芽雌蕊分化期叶片可溶性糖与淀粉含量显著低于其他时期;叶片P、K、Ca含量在分化前期较高,但是雌蕊、雄蕊分化期显著下降。(4)与正常花相比,败育花主要表现为子房发育异常,多数生长点不均匀,子房萎缩,胚珠原基发育停滞;雌蕊的花柱低于花丝,且花粉粒急剧减少。研究发现,‘李光杏’树势的强弱会造成败育花比例和坐果率的差异,并以中庸树果实坐果率最高,且花败育率最低;雌蕊发育是否正常直接关系到‘李光杏’正常开花,雌蕊的发育受阻,最终成为整个花芽退化的主要形式;叶片碳水化合物和矿质元素等营养物质参与并保证了花器官的正常发育。     

In Loquat (Eriobotrya japonica Lindl.) Return Bloom Depends on the Time the Fruit Remains on the Tree

In loquat (Eriobotrya japonica Lindl.), the comparison of fruiting trees and defruited trees carried out covering a range of developmental fruit stages reveals a significant reduction in flowering due to fruit from its early stage of development, being higher when it changes color and becomes senescent, which coincides with the floral bud inductive period. This effect occurred both at the tree and at the shoot level. Furthermore, although current shoots almost always develop into panicles, those from fruiting trees develop fewer flowers, suggesting that fruit also affects at the floral bud level. In our experiment, the gibberellin concentration at the floral bud inductive period was significantly higher in bark tissues (periderm, cortex and phloem tissues) of fruiting trees, compared with defruited trees that tend to flower more. The lower concentration of IAA in the bark tissues of defruited trees also contributes to increase their flowering intensity. On the contrary, the zeatin concentration was higher. Accordingly, at bud burst, the IAA/zeatin ratio, an indication of effect on flowering, was significantly higher for fruiting trees. Some disruption in the nitrate reduction process in fruiting tree was also observed. The process of floral bud induction and differentiation was not associated with either reducing or translocating and reserve carbohydrate concentration. Hence, loquat flower intensity depends on the time the fruit is maintained on the tree. The intensity is affected indirectly, by reducing the number of shoots, and directly, by reducing the number of flowers per panicle, and these effects are linked to endogenous plant hormone contents.     

Factors affecting pollinator movement and plant fitness in a specialized pollination system

The rate of pollen exchange within and among flowers may depend on pollinator attraction traits such as floral display size and flowering plant density. Variations in these traits may influence pollinator movements, pollen receipt, and seed number. To assess how floral display size and flowering plant density affect parameters of pollinator visitation rate, pollen receipt per flower, seed number per fruit and the between-plant pollinator movements, we studied the self-incompatible plant, Nierembergia linariifolia. Per-flower pollinator visitation rate and bout length increased linearly with increasing floral display size. Pollen receipt per flower increased linearly with increasing flowering plant density. For seed number per fruit, a polynomial model describing an increased seed number per fruit at low density and a decreased seed number per fruit at high density provided a significant fit. Per-flower pollinator visitation rate was not associated with pollen receipt per flower and seed number per fruit. Bees visited plants located near to the center of the population more frequently than plants located at the periphery. Increases in both floral display size and flowering plant density led to an increased chance of a plant being chosen as the center of the pollinator foraging area. These results suggest that even though large floral displays and high flowering plant density are traits that attract more pollinators, they may also reduce potential mate diversity by restricting pollen movement to conspecific mates that are closely located.     

油茶花芽分化形态结构及内源激素变化

以普通油茶(Camellia oleifera) 4个无性系为材料,结合油茶成花的动态观察和花芽分化过程的石蜡切片形态观察,采用酶联免疫吸附分析法测定花芽中玉米素核苷(ZR)、脱落酸(ABA)、生长素(IAA)、赤霉素(GA) 4种内源激素含量,探讨油茶花芽分化与内源激素的关系。油茶花芽分化过程可分为6个时期：前分化期(10 d)、萼片形成期(20 d)、花瓣形成期(30 d)、雌雄蕊形成期(20 d)、子房与花药形成期(10 d)和雌雄蕊成熟期(20 d),历时3～4个月。油茶不同无性系的花芽分化时间略有不同。油茶花芽中ZR含量相对较低(5.102～16.412 ng·g–1 FW),ABA含量相对较高(76.815～137.648 ng·g–1 FW)。其中,粤华5号和湘林8号的ZR、ABA含量变化趋势一致,岑软3号和岑软2号含量变化趋势一致。油茶花芽中IAA含量相对较高,为49.072～135.622 ng·g–1 FW,随着花芽分化进程,IAA含量均呈先升后降再升的变化趋势。GA含量相对较低,为5.616～13.720 ng·g–1 FW,随时间变化,呈现出不断降低的趋势。其中,不同无性系的IAA、GA含量变化趋势一致,而ZR、ABA含量变化趋势有所差异。ZR有利于花器官形成;高浓度IAA促进油茶花芽分化,低浓度IAA有利于开花;花芽中IAA与ABA存在明显的颉颃作用;GA抑制花芽分化。     

Gibberellic Acid Reduces Flowering Intensity in Sweet Orange [Citrus sinensis (L.) Osbeck] by Repressing CiFT Gene Expression

In Citrus, gibberellic acid (GA₃) applied at the floral bud inductive period significantly reduces flowering intensity. This effect is being used to improve the fruit set of parthenocarpic cultivars that tend to flower profusely. However, the molecular mechanisms involved in the process remain unclear. To contribute to the knowledge of this phenomenon, adult trees of ‘Salustiana’ sweet orange were sprayed at the floral bud inductive period with 40?mg?L^?1 of GA₃ and the expression pattern of flowering genes was examined up to the onset of bud sprouting. Trees sprayed with paclobutrazol (PBZ, 2,000?mg L^?1), a gibberellin biosynthesis inhibitor, were used to confirm the effects, and untreated trees served as control. Bud sprouting, flowering intensity, and developed shoots were evaluated in the spring. GA₃ significantly reduced the number of flowers per 100 nodes by 72% compared to the control, whereas PBZ increased the number by 123%. Data of the expression pattern of flowering genes in leaves of GA₃-treated trees revealed that this plant growth regulator inhibited flowering by repressing relative expression of the homolog of FLOWERING LOCUS T, CiFT, whereas PBZ increased flowering by boosting its expression. The activity of the homologs TERMINAL FLOWER 1, FLOWERING LOCUS C, SUPPRESSOR OF OVEREXPRESSION OF CONSTANS 1, and APETALA1 was not affected by the treatments. The number of flowers per inflorescence, in both leafy and leafless inflorescences, was not altered by GA₃ but increased with PBZ; the latter paralleled LEAFY relative expression. These results suggest that GA₃ inhibits flowering in Citrus by repressing CiFT expression in leaves.     

施硼和赤霉素对‘李广杏’坐果率及果实品质的影响

该研究以8年生盛果期的‘李广杏’植株为试材,于花蕾膨大期喷施0.1%(P1)、0.3%(P2)和0.5%(P3)硼砂和盛花初期喷施50(C1)、100(C2)和150 mg/L(C3)赤霉素,测定不同处理下李广杏的坐果率、果实品质及营养生长指标的变化,并用主成分分析不同处理的效果进行综合评价,为敦煌寒旱区‘李广杏’的栽培提供理论依据。结果表明:(1)‘李广杏’叶面积增长量在C1浓度处理下显著高于CK,不同浓度的硼处理对其均具有抑制作用,但C1、P2浓度下新稍生长量高于其他处理。(2)适宜浓度的硼和赤霉素处理可一定程度减少‘李广杏’花的败育率,从而有效提高果树的坐果率,其中硼处理以P3浓度下最优,但P3与P2处理下坐果率无显著差异,赤霉素处理以C2浓度下最优。(3)适宜浓度硼和赤霉素能够明显提高‘李广杏’果实品质,C1浓度处理下的果实糖酸比、可溶性固形物含量显著高于CK,P2处理下的果实维生素C、可滴定酸含量显著高于其他处理;适宜浓度赤霉素和硼对果实的单果重、果形指数、侧径有明显的促进作用。(4)主成分分析结果显示,各处理效果的综合得分由高到低依次为P2(1.20)>C2(0.91)>P1(0.13)>C1(-1.01)>CK(-1.68)>P3(-2.13)>C3(-7.76);果实可溶性糖、可溶性蛋白、糖酸比等主成分占比较高,可作为评价‘李广杏’果实品质的重要依据。研究发现,花期喷施适宜浓度的硼和赤霉素可有效提高‘李广杏’果树的生长状况、坐果率及果实品质,且甘肃敦煌地区以花蕾膨大期喷施0.3%硼和盛花初期喷施100 mg/L赤霉素效果最佳。     

穗花牡荆花芽分化过程中形态和生理指标变化

以穗花牡荆为研究材料,通过探究其花芽分化进程和生理特性,为花期调控技术提供成花机理。采用物候期观察和石蜡切片相结合的方法并测定花芽分化过程中相关生理指标,研究花发育过程中的形态和生理变化。结果表明,穗花牡荆花芽分化为一年多次分化型,其进程可划分为七个时期：未分化期、总轴花序原基分化期、初级分轴花序原基分化期、次级分轴花序原基分化期、小花原基分化期、花器官分化前期和花器官分化后期。同一植株不同位置花芽及同一花序中不同单花分化的进程不同,第一季花期后各阶段的花芽分化形态常存在重叠。花芽分化过程中不同时期叶片和花芽的可溶性糖和可溶性蛋白质含量均有上升下降的变化,总体上叶片中营养物质含量高于花芽保证营养供应。花芽分化过程中,IAA、ABA、CTK和GA3整体水平上先升后降有利于花芽分化进行。研究认为,花芽中大量的可溶性糖和蛋白质积累及较高的碳氮比,有利于穗花牡荆花芽形态分化顺利完成。低水平的GA3/ABA和IAA/CTK有利于花序的形成,ABA/CTK和ABA/IAA比值升高促进小花原基和小花萼片原基的分化, GA3/CTK、GA3/ABA和GA3/IAA比值升高促进花瓣原基、雄雌蕊原基发育。     

Sex‐specific phenotypic selection and geographic variation in gender divergence in a gynodioecious shrub

In sexually polymorphic plant species the extent of gender divergence in floral morphology and phenology may be influenced by gender‐specific selection patterns imposed by pollinators, which may change geographically. Distribution margins are areas where changes in the pollinator fauna, and thus variation in gender divergence of floral traits, are expected. We tested for pollination‐driven geographic variation in the gender divergence in floral and phenological traits in the gynodioecious shrub Daphne laureola, in core and marginal areas differing in the identity of the main pollinator. Pollinators selected for longer corolla tubes in hermaphrodite individuals only in core populations, which in turn recorded higher fruit set. Consistent with these phenotypic selection patterns, gender divergence in flower corolla length was higher in core populations. Moreover, pollinators selected towards delayed flowering on hermaphrodite individuals only in marginal populations, where the two sexes differed more in flowering time. Our results support that a shift in main pollinators is able to contribute to geographic variation in the gender divergence of sexually polymorphic plant species.     

Reproductive biology of the epiphytic bromeliad Werauhia gladioliflora in a premontane tropical forest

The floral phenology, fruit and seed production, and self-compatibility of Werauhia gladioliflora, an epiphytic bromeliad with a wide distribution, were studied in a premontane forest in the Monteverde area in Costa Rica. The species presents the pollination syndrome of chiropterophily, and it is visited by the small bats Hylonycteris underwoodi and Glossophaga commissarisi (Glossophaginae). The population flowering period extended from October to early December (end of rainy season) and seed dispersal occurred from February to April (dry season). Most plants opened a single flower per night, either every day or at one-day intervals during the flowering period. In natural conditions, the average fruit set amounted to almost half of the potential output, but individual fecundity (number of seeds) remained high. Seed number per fruit and germination capacity after artificial selfing and out-crossing treatments did not differ from natural pollination conditions. Werauhia gladioliflora exhibited high levels of autonomous self-pollination and self-compatibility at the individual and population level, characters associated with the epiphytic habitat. These reproductive traits are also associated with early colonizer species, yet life history traits, such as seed dispersal, seedling establishment success, and growth, are likely to have a major role in determining the presence of this species in the successional vegetation patches scattered over the studied premontane area.     

Genetic variation in the reduction of attractive floral traits of an annual tarweed in response to drought and apical damage

Aims Foliar herbivory and water stress may affect floral traits attractive to pollinators. Plant genotypes may differ in their responses to the interplay between these factors, and evolution of phenotypic plasticity could be expected, particularly in heterogeneous environments. We aimed at evaluating the effects of simulated herbivory and experimental drought on floral traits attractive to pollinators in genetic families of the annual tarweed Madia sativa, which inhabits heterogeneous environments in terms of water availability, herbivore abundance and pollinator abundance.Methods In a greenhouse experiment with 15 inbred lines from a M. sativa population located in central Chile (Mediterranean-type climate), we measured the effects of apical bud damage and reduced water availability on: number of ray florets per flower head, length of ray florets, flower head diameter, number of open flower heads per plant, flowering plant height and flowering time.Important findings Apical damage and water shortage reduced phenotypic expression of floral traits attractive to pollinators via additive and non-additive effects. Plants in low water showed decreased height and had fewer and shorter ray florets, and fewer and smaller flower heads. Damaged plants showed delayed flowering, were less tall, and showed shorter ray florets and smaller flower heads. The number of ray florets was reduced by damage only in the low water treatment. Plant height, flowering time and number of flower heads showed among-family variation. These traits also showed genetic variation for plasticity to water availability. Ray floret length, flower head size and time to flowering showed genetic variation for plastic responses to apical damage. Plasticity in flowering time may allow M. sativa to adjust to the increased aridity foreseen for its habitat. Because genetic variation for plastic responses was detected, conditions are given for evolutionary responses to selective forces acting on plastic traits. We suggest that the evolution of adaptive floral plasticity in M. sativa in this ecological scenario (heterogeneous environments) would result from selective forces that include not only pollinators but also resource availability and herbivore damage.     

高等植物开花结实的多胺研究进展

多胺是广泛分布于植物体内具有调控作用的生理活性物质,其代谢变化与高等植物的 生长和发育关系密切。本文概述了多胺与植物花芽形成、花器官分化以及开花、坐果和果实发育的关系;并就外源多胺对植物开花坐果的影响做了述评,对多胺在植物开花结实中可能 的作用机理及今后的研究方向和应用潜力进行了讨论。     

高等植物开花结实的多胺研究进展

多胺是广泛分布于植物体内具有调控作用的生理活性物质,其代谢变化与高等植物的生长和发育关系密切。本文概述了多胺与植物花芽形成、花器官分化以及开花、坐果和果实发育的关系;并就外源多胺对植物开花坐果的影响做了述评,对多胺在植物开花结实中可能的作用机理及今后的研究方向和应用潜力进行了讨论。     

A Developmental Pattern of Flowering in Colored <Emphasis Type="Italic">Zantedeschia</Emphasis> spp.: Effects of Bud Position and Gibberellin

The restricted flowering of colored cultivars ofZantedeschia is a consequence of developmental constraints imposed by apical dominance of the primary bud on secondary buds in the tuber, and by the sympodial growth of individual shoots. GA₃ enhances flowering inZantedeschia by increasing the number of flowering shoots per tuber and inflorescences per shoot. The effects of gibberellin on the pattern of flowering and on the developmental fate of differentiated inflorescences along the tuber axis and individual shoot axes were studied in GA₃ and Uniconazole-treated tubers. Inflorescence primordia and fully developed (emerged) floral stems produced during tuber storage and the plant growth period were recorded. Days to flowering, percent of flowering shoots and floral stem length decreased basipetally along the shoot and tuber axes. GA₃ prolonged the flowering period and increased both the number of flowering shoots per tuber and the differentiated inflorescences per shoot. Activated buds were GA₃ responsive regardless of meristem size or age. Uniconazole did not inhibit inflorescence differentiation but inhibited floral stem elongation. The results suggest that GA₃ has a dual action in the flowering process: induction of inflorescence differentiation and promotion of floral stem elongation. The flowering pattern could be a result of a gradient in the distribution of endogenous factors involved in inflorescence differentialtion (possibly GAs) and in floral stem growth. This gradient along the tuber and shoot axes is probably controlled by apical dominance of the primary bud. Online publication: 7 April 2005