植物成花转变及成花逆转的研究进展

本文主要讨论了植物成花转变及成花逆转等开花研究中应用分子生物学技术所取得的一些进展,描述了成花决定态的特性,5个从拟南芥中分离获得的成花转变的基因,有关光受体的分子生物学研究和造成成花逆转的原因等。     

植物开花时间调控的信号途径

开花是植物从营养生长到生殖生长的一个重要转折点。花启动的时机对生殖生长的成功至关重要。开花时间受内在因子和环境因子的共同调节。通过对拟南芥的分子遗传学研究,确定至少存在4条调控开花时间的信号途径,即光周期途径、春化途径、自主途径和赤霉素途径。本文以拟南芥 (Arabidopsis thaliana) 为主要研究对象简要综述了近年来在开花时间调控领域的研究进展。     

Indirect effects of FRIGIDA: floral trait (co)variances are altered by seasonally variable abiotic factors associated with flowering time

Reproductive timing is a critical life‐history event that could influence the (co)variation of traits developing later in ontogeny by regulating exposure to seasonally variable factors. In a field experiment with Arabidopsis thaliana, we explore whether allelic variation at a flowering‐time gene of major effect (FRIGIDA) affects (co)variation of floral traits by regulating exposure to photoperiod, temperature, and moisture levels. We detect a positive latitudinal cline in floral organ size among plants with putatively functional FRI alleles. Statistically controlling for bolting day removes the cline, suggesting that seasonal abiotic variation affects floral morphology. Both photoperiod and precipitation at bolting correlate positively with the length of petals, stamens, and pistils. Additionally, floral (co)variances differ significantly across FRI backgrounds, such that the sign of some floral‐trait correlations reverses. Subsequent experimental manipulations of photoperiod and water availability demonstrate direct effects of these abiotic factors on floral traits. In sum, these results highlight how the timing of life‐history events can affect the expression of traits developing later in ontogeny, and provide some of the first empirical evidence for the effects of major genes on evolutionary potential.     

长角凤仙花（凤仙花科）的花器官发生和发育

以不同发育时期的长角凤仙花Impatiens longicornuta Y.L.Chen（凤仙花科Balsaminaceae）为材料,利用扫描电镜技术观察了其花器官的分化及其发育过程。长角凤仙花为两侧对称花,具2枚侧生萼片,唇瓣囊状,旗瓣具鸡冠状突起,雄蕊5枚,子房上位,5心皮5室。其花器官分化顺序为向心式,萼片—花瓣—雄蕊—雌蕊原基。2枚侧生萼片先发生,然后近轴萼片（即唇瓣）原基和2枚前外侧萼片原基近同时发生;但是这3枚萼片原基的发育不同步,远轴的2枚前外侧萼片原基的发育渐渐滞后,然后停止发育,最后渐渐为周围组织所吸收,直至消失不见。花瓣原基中,旗瓣原基最先发生,4个侧生花瓣原基相继成对发生,且之后在基部成对愈合形成翼瓣;5枚雄蕊原基几乎同时发生,5个心皮原基轮状同时发生。本文结果支持凤仙花属植物为5基数的花,并进一步证实了唇瓣的萼片来源;此外,研究结果表明花器官早期发育资料对植物系统与进化研究具有重要参考价值。     

Florigen goes molecular: Seventy years of the hormonal theory of flowering regulation

As early as in 1936, the comprehensive studies of flowering led M.Kh. Chailakhyan to the concept of florigen, a hormonal floral stimulus, and let him establish several characteristics of this stimulus. These studies set up for many years the main avenues for research into the processes that control plant flowering, and the notion of florigen became universally accepted by scientists worldwide. The present-day evidence of genetic control of plant flowering supports the idea that florigen participates in floral signal transduction. The recent study of arabidopsis plants led the authors to conclusion that the immediate products of the gene FLOWERING LOCUS I, its mRNA and/or protein, move from an induced leaf into the shoot apex and evoke flowering therein.     

Experimental control of floral reversion in isolated shoot apices of the long-day plant Silene coeli-rosa

The fate of shoot meristems of the long day (LD) plant Silene coeli-rosa in culture was examined (complete, reverted or arrested flowers) to establish whether these different patterns were related to a particular stage of morphogenesis and the extent to which the fate of the pattern was regulated by either added plant growth regulators (PGRs) or changing the carbohydrate source in the medium. In particular, the frequency of reversion was measured to test the stability of the determined state for each whorl. The plants were given various inductive treatments (4–7 LD, 7 LD + 1 to 3 SD) and the apices were explained onto Murashige-Skoog medium supplemented with 3% sucrose (controls) ± IAA, ± kinetin, ± GA₃ or onto the basal medium containing 1 or 3% sucrose, glucose or fructose or 7% sucrose. The apices were examined 12 weeks later. When the data were pooled from all inductive treatments, IAA resulted in more reversions, GA₃ caused more arrested flowers while kinetin hardly affected the pattern of meristem fate compared with the controls. However, each PGR treatment did not perturb the pattern of organ formation for those apices that formed either arrested or complete flowers. The time for determination (days) of the earlier formed whorls (determination times for the controls in brackets): sepals (2), stamens 1–5 (3) and petals (3), was shortened by about a day in all PGR treatments whereas the corresponding times for the later formed whorls: stamens 6–10 (4) and carpels (4), were either lengthened to 5 days or unaffected. The response of the apices to the various sugars was simply a reflection of concentration. Hence, more complete flowers formed at 7 or 3% and more flowers were arrested at 1 % regardless of the sugar moiety. However, the frequency of reversion was similar on each of the media. Pooling all data from all treatments enabled a statistical analysis of the pattern of reversion and the pattern of arrest. Reversion was more common from apices which exhibited the later-formed whorls (stamens 6–10 and carpels) than from the earlier whorls. Moreover, the stronger the inductive treatment the less frequent was reversion. The most common stage of arrest was at the stamen 6–10 whorl and this was particularly so for the GA₃ treatment. The data indicated that reversion could occur from any whorl, which suggests that determination of each whorl is independent of the next. This conclusion is underlined by the more frequent occurrence of reversion from the carpel whorl. However, the longer the inductive treatment the less likelihood of reversion; this suggests that in Silene, the floral stimulus is required continuously to stabilise the determined state of each whorl and to ensure smooth completion of floral morphogenesis.     

拟南芥开花时间调控的分子基础

在合适的时间开花对大多数植物的生存和成功繁衍极为重要。开花时间受错综复杂的环境因素和植物自身的遗传因子影响,由开花调控因子所构成的光周期、春化、温度、赤霉素、自主以及年龄等至少6条既相互独立又相互联系的遗传途径调控。该文综述了有关拟南芥(Arabidopsis thaliana)开花时间调控的分子机制的最新研究进展,并对今后的研究进行了展望。     

Selection on flowering time and floral display in an alpine and a lowland population of Arabidopsis lyrata

To determine whether population differentiation in flowering time is consistent with differences in current selection, we quantified phenotypic selection acting through female reproductive success on flowering phenology and floral display in two Scandinavian populations of the outcrossing, perennial herb Arabidopsis lyrata in two years. One population was located in an alpine environment strongly affected by grazing, whereas the other was close to sea level and only moderately affected by herbivory. Multiple regression models indicated directional selection for early end of flowering in one year in the lowland population, and directional selection for early start of flowering in one year in the alpine population. As expected, there was selection for more inflorescences in the lowland population. However, in the alpine population, plants with many inflorescences were selectively grazed and the number of inflorescences produced was negatively related to female fitness in one year and not significantly related to female fitness in the second year. The results are consistent with the hypothesis that genetic differentiation in flowering phenology between the study populations is adaptive, and indicate that interactions with selective grazers may strongly influence selection on floral display in A. lyrata.     

拟南芥开花时间调控的研究进展

调控开花时间是大多数植物由营养生长向生殖生长转化的一个重要生长发育过程.影响拟南芥开花时间的因素有很多,其中光照和温度是两个主要的外部因素,而赤霉素(GA)和一些自主性因子是主要的内部因素.目前,一般按照对以上因素的反应将晚花突变体归于四条开花调控途径:光周期途径、春化途径、自主途径和GA途径.在不断变化的外部环境条件和内部生理条件下,这些途径通过一些主要的整合基因如SOC1、FT、LFY等实现了对拟南芥开花时间的精确调控.     

A comparison of in vitro with in vivo flowering in bamboo: Bambusa arundinacea

Seedling explants of Bambusa arundinacea were cultured in a Murashige & Skoog (MS) based liquid medium, supplemented with sucrose (2), coconut water (5) and 6-benzylaminopurine (2.2 μM). In 3–6 months about 70 of the cultures flowered. A comparison was made between in vitro and in vivo flowering. Though smaller in size, in vitro florets were morphologically comparable to the in vivo florets. Anthesis in in vivo flowering took place in the morning hours. It was more or less synchronized and was dependent on the atmospheric temperature and humidity. The lemma and palea opened to expose both androecium and gynoecium to the pollinating agent (wind). In in vitro flowering, some florets opened as in their in vivo counterparts, some did not open but the anthers protruded from the tip of the partially opened lemma and palea. Anthesis was not synchronized under in vitro conditions. Pollen fertility in in vivo and in vitro flowerings were approximately 93 and 31 respectively. Studies by scanning electron microscopy showed some discrepancies in the pollen wall development in vitro. The trifid stigmas of in vivo florets were highly feathery with many papillae and withered soon after pollination or within few hours. The stigmas of in vitro developed florets were smaller with fewer and stouter papillae. They remained turgid for relatively longer periods. Seed production in in vivo flowering was profuse whereas in in vitro flowering seeds were produced only when many florets opened at the same time, in the same culture vessel. This revised version was published online in June 2006 with corrections to the Cover Date.     

APETALA1启动子驱动AtIPT4在转基因拟南芥中表达导致花和花器官发育异常

细胞分裂素对拟南芥（Arabidopsis thaliana）花分生组织细胞的分裂和分化具有重要作用。本研究利用APETALA1（AP1）特异启动子在花分生组织和第1、2轮花器官中表达细胞分裂素合成酶（isopentyl transferase,IPT）基因IPT4,研究细胞分裂素对花和花器官发育的影响。在pAP1∷IPT4转基因植株中出现了花密集和花器官数目增多等现象。原位杂交和GUS组织染色结果发现,在pAP1∷IPT4转基因植株中,花分生组织特征决定基因LEAFY（LFY）与花器官特征决定基因AP1、PISTILLATA（PI）和AGAMOUS（AG）的表达量均有不同程度的提高。研究结果表明在拟南芥中表达pAP1∷IPT4影响其花和花器官的正常发育。     

高等植物成花分子机理研究现状及展望

以拟南芥、金鱼草为例,介绍了近几年植物成花（包括成花诱导、花序分生组织的组成、花发端、花器官发生及发育）研究的一些进展,着重介绍了成花过程中基因的表达,调控与生理功能。     

华凤仙花部特征和传粉系统研究

研究了华凤仙（Impatiens chinensis L.）的花部特征和传粉系统,结果表明华凤仙单花期为2.3 d,雄性期是雌性期的6.8倍。花蜜量和花蜜可溶性糖含量分别为0.56 μL和28.2%。开花当天柱头即具有可授性,花粉活力在花朵刚开放时最高（95%）,至花朵凋谢之前仍保持较高的活力（55%）。花粉胚珠比（5 730.3±2 941.1）和套袋实验的结果表明华凤仙属于自交亲和但以异交为主的繁育系统,传粉过程需要传粉者。蜜蜂、熊蜂、蝶类和几种芦蜂是华凤仙的主要访花者,最有效的传粉者为三条熊蜂。考氏无垫蜂作为盗蜜者,可能是除正常访花者之外,对华凤仙的有性繁殖影响最大的生物因素。