高等植物开花时程的基因调控（Ⅰ）

高等植物从营养生长向生殖生长及发育转变的时程具有重要意义,但是了解得很少。近6年来利用分子遗传学方法详细地分析了拟南芥中的这一转变的时程变化,为高等植物开花时程的基因调控提供了一个很好的模式。有关早期或晚期开花表现型的大量突变体及遗传变异得到了阐述。这里谈到的表现型对影响开花转变的环境及内部因子的控制有重大作用。通过分子生物学、遗传学和生理学分析已经鉴定了参与此过程的不同组分,如光识别和昼夜节律（circadian rhythm）因子。另外,通过克隆某些花诱导基因及其相应的靶基因已经对参与开花信号转导途径（signal transduction pathway）的相关因子进行了系统的鉴定,这些开创性工作大大促进了高等植物开花时程的基因表达调控研究及其机理的阐明。本实验室在以黄瓜、新红宝西瓜、西葫芦为材料所获得的部分结果基础上,主要以近六年来在拟南芥方面获得的进展为依据,对高等植物开花时程的基因调控作一系统的总结,并对其开花时程基因调控的机理提出可能的作用理论模型。     

高等植物开花时程的调控与光受体Ⅰ.开花时程的基因与光受体调控

系统评述了高等植物开花时程的调控与植物光受体的联系.重点说明了控制开花时程的遗传途径以及光周期途径的有关基因的研究进展.影响高等植物开花的最重要的因子之一便是光周期,光周期对高等植物开花的调控是通过相关基因间的相互作用来实现的,这些基因包括参与花启动发育控制基因,昼夜节律时间钟调控基因及光受体信号转导基因.近5年左右的时间通过对拟南芥及其一系列突变体的研究为我们展示了这一热门领域的广阔的前景.     

模式植物拟南芥开花时间分子调控研究进展

植物从营养生长到生殖生长的转变是开花发育的关键,在合适的时间开花对植物的生长和繁衍极为重要,植物开花时间的调控对农业生产发展意义重大。植物开花是由遗传因子和环境因子协同调节的一个复杂过程。近年来,对不同植物开花调控的研究,特别是对模式植物拟南芥（Arabidopsis thaliana（L.） Heynh.）的开花调控研究取得了显著进展,已探明开花时间分子调控的6条主要途径分别是光周期途径、春化途径、自主途径、温度途径、赤霉素途径和年龄途径。各遗传调控途径既相互独立又相互联系,构成一个复杂的开花调控网络。本文综述了模式植物拟南芥开花时间调控分子机制相关研究的最新进展,并对未来的研究进行了展望。     

拟南芥开花过程的表观遗传调节

开花是高等植物由营养生长到生殖生长的重要转变.拟南芥的开花时间受许多基因的调控,其中一些基因的表现遗传调节对开花时间的控制发挥着重要的作用.本文就这些基因以表观遗传方式调节拟南芥开花过程的最新研究进程作简要介绍.     

高等植物开花诱导途径信号整合的分子机制

开花是高等植物从营养生长到生殖生长的重要转折点。花分生组织的形成是开花植物对内外环境信号的响应。近年来在开花诱导方面已获得许多研究成果,我们介绍了高等植物开花诱导的4条主要途径（光周期途径、春化途径、自主途径和赤霉素途径）和复杂的信号整合机制。     

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。     

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.     

高等植物开花时程的调控与光受体Ⅱ.光受体调控开花时程的分子机制与昼夜节律时间钟

系统评述了高等植物开花时程的调控与植物光受体的联系.重点说明了控制开花时程的遗传途径以及光周期途径的有关基因的研究进展.而且对植物光受体调控高等植物开花里程的分子机制作了深入的探讨.高等植物从营养生长向生殖生长及发育转变的时程具有重要意义.控制高等植物开花时程及其性别表达的关键就在此过程中.植物光受体参与了高等植物开花时程的调控并起到了重要作用.植物光受体主要包括植物光敏素受体(光敏素A、B、D、E受体)和隐色素受体.近5年左右的时间通过对拟南芥及其一系列突变体的研究展示了这一热门领域的广阔的理论与应用前景.     

MADS-box基因控制植物成花的分子机理

植物花器官的发育和开花是植物生殖发育中最重要的过程,植物在长期的进化过程中产生了春化(低温)途径、自主途径、光周期途径以及不依赖于光温环境条件的赤霉素信号途径来适应多变的环境和调控植物开花过程。本文综述了模式植物拟南芥中由LEAFY(LFY)、CONSTANS(CO)、FLOWERING LOCUSC(FLC)、FLOW ERING LOCUS T(FT)和SUPPRESSOR OF OVEREXPRESSION OF CO1(SOC1)等基因构成的双子叶植物响应光温条件变化的开花调控网络;以及大麦、小麦中由VERNALIZATION1(VRN1)、VRN2、ODD-SOC2(OS2)和拟南芥CO、FT同源基因构成的禾本科植物开花调控网络。其中最重要的是转录调控因子MADS-box基因FLC、SOC1、VRN1和OS2,并发现组蛋白的乙酰化/脱乙酰化,赖氨酸的甲基化/脱甲基化在调控FLC、VRN1染色质活性状态及基因表达,从而产生开花控制的机理。这些研究发现将有助于对具有重要经济价值的单双子叶植物,通过生物技术手段改良其品种特性以应对非生物逆境,特别是低温胁迫的指导。     

拟南芥开花诱导途径分子机制研究进展

拟南芥是分子和遗传学研究的模式植物,对植物花发育及控制花形态建成的分子遗传机制的研究进展主要是建立在对拟南芥研究的基础之上,拟南芥开花主要受到4个途径(自主途径、赤霉素途径、春化作用和光周期途径)的内源和外界信号的同时诱导.该文对近年来国内外有关拟南芥开花诱导的4个途径的分子机制研究进展进行综述,并初步绘制出各开花诱导途径基因间的调控网络图,以进一步明确基因间的相互作用模式及其在整个开花过程中的作用地位.     

New insights into gibberellin signaling in regulating flowering in Arabidopsis

In angiosperms,floral transition is a key developmental transition from the vegetative to reproductive growth,and requires precise regulation to maximize the reproductive success.A complex regulatory network governs this transition through integrating flowering pathways in response to multiple exogenous and endogenous cues.Phytohormones are essential for proper plant developmental regulation and have been extensively studied for their involvement in the floral transition.Among various phytohormones,gibberellin(GA)plays a major role in affecting flowering in the model plant Arabidopsis thaliana.The GA pathway interact with other flowering genetic pathways and phytohormone signaling pathways through either DELLA proteins or mediating GA homeostasis.In this review,we summarize the recent advances in understanding the mechanisms of DELLA-mediated GA pathway in flowering time control in Arabidopsis,and discuss its possible link with other phytohormone pathways during the floral transition.     

Wheat SOC1 functions independently of WAP1/VRN1, an integrator of vernalization and photoperiod flowering promotion pathways

Heading time in bread wheat ( Triticum aestivum L.) is determined by three characters – vernalization requirement, photoperiodic sensitivity and narrow-sense earliness (earliness per se) – which are involved in the phase transition from vegetative to reproductive growth. The wheat APETALA1 ( AP1 )-like MADS-box gene, wheat AP1 ( WAP1 , identical with VRN1 ), has been identified as an integrator of vernalization and photoperiod flowering promotion pathways. A MADS-box gene, SUPPRESSOR OF OVEREXPRESSION OF CO 1 ( SOC1 ) is an integrator of flowering pathways in Arabidopsis . In this study, we isolated a wheat ortholog of SOC1 , wheat SOC1 ( WSOC1 ), and investigated its relationship to WAP1 in the flowering pathway. WSOC1 is expressed in young spikes but preferentially expressed in leaves. Expression starts before the phase transition and is maintained during the reproductive growth phase. Overexpression of WSOC1 in transgenic Arabidopsis plants caused early flowering under short-day conditions, suggesting that WSOC1 functions as a flowering activator in Arabidopsis . WSOC1 expression is affected neither by vernalization nor photoperiod, whereas it is induced by gibberellin at the seedling stage. Furthermore, WSOC1 is expressed in transgenic wheat plants in which WAP1 expression is cosuppressed. These findings indicate that WSOC1 acts in a pathway different from the WAP1 -related vernalization and photoperiod pathways.     

A MADS domain gene involved in the transition to flowering in Arabidopsis

Flowering time in many plants is triggered by environmental factors that lead to uniform flowering in plant populations, ensuring higher reproductive success. So far, several genes have been identified that are involved in flowering time control. AGL20 (AGAMOUS LIKE 20) is a MADS domain gene from Arabidopsis that is activated in shoot apical meristems during the transition to flowering. By transposon tagging we have identified late flowering agl20 mutants, showing that AGL20 is involved in flowering time control. In previously described late flowering mutants of the long-day and constitutive pathways of floral induction the expression of AGL20 is down-regulated, demonstrating that AGL20 acts downstream to the mutated genes. Moreover, we can show that AGL20 is also regulated by the gibberellin (GA) pathway, indicating that AGL20 integrates signals of different pathways of floral induction and might be a central component for the induction of flowering. In addition, the constitutive expression of AGL20 in Arabidopsis is sufficient for photoperiod independent flowering and the over-expression of the orthologous gene from mustard, MADSA, in the classical short-day tobacco Maryland Mammoth bypasses the strict photoperiodic control of flowering.     

Flowering responses to light and temperature

Light and temperature signals are the most important environmental cues regulating plant growth and development. Plants have evolved various strategies to prepare for, and adapt to environmental changes. Plants integrate environmental cues with endogenous signals to regulate various physiological processes, including flowering time. There are at least five distinct pathways controlling flowering in the model plant Arabidopsis thaliana: the photoperiod pathway, the vernalization/thermosensory pathway, the autonomous floral initiation, the gibberellins pathway, and the age pathway. The photoperiod and temperature/vernalization pathways mainly perceive external signals from the environment, while the autonomous and age pathways transmit endogenous cues within plants. In many plant species, floral transition is precisely controlled by light signals(photoperiod) and temperature to optimize seed production in specific environments. The molecular mechanisms by which light and temperature control flowering responses have been revealed using forward and reverse genetic approaches. Here we focus on the recent advances in research on flowering responses to light and temperature.     

What is going on with the hormonal control of flowering in plants?

Molecular genetic studies using Arabidopsis thaliana as a model system have overwhelmingly revealed many important molecular mechanisms underlying the control of various biological events, including floral induction in plants. The major genetic pathways of flowering have been characterized in-depth, and include the photoperiod, vernalization, autonomous and gibberellin pathways. In recent years, novel flowering pathways are increasingly being identified. These include age, thermosensory, sugar, stress and hormonal signals to control floral transition. Among them, hormonal control of flowering except the gibberellin pathway is not formally considered a major flowering pathway per se, due to relatively weak and often pleiotropic genetic effects, complex phenotypic variations, including some controversial ones. However, a number of recent studies have suggested that various stress signals may be mediated by hormonal regulation of flowering. In view of molecular diversity in plant kingdoms, this review begins with an assessment of photoperiodic flowering, not in A. thaliana, but in rice (Oryza sativa); rice is a staple crop for human consumption worldwide, and is a model system of short-day plants, cereals and breeding crops. The rice flowering pathway is then compared with that of A. thaliana. This review then aims to update our knowledge on hormonal control of flowering, and integrate it into the entire flowering gene network.     

The role of auxin and sugar signaling in dominance inhibition of inflorescence growth by fruit load

Dominance inhibition of shoot growth by fruit load is a major factor that regulates shoot architecture and limits yield in agriculture and horticulture crops. In annual plants, the inhibition of inflorescence growth by fruit load occurs at a late stage of inflorescence development termed the end of flowering transition. Physiological studies show this transition is mediated by production and export of auxin from developing fruits in close proximity to the inflorescence apex. In the meristem, cessation of inflorescence growth is controlled in part by the age-dependent pathway, which regulates the timing of arrest. Here, we show the end of flowering transition is a two-step process in Arabidopsis (Arabidopsis thaliana). The first stage is characterized by a cessation of inflorescence growth, while immature fruit continues to develop. At this stage, dominance inhibition of inflorescence growth by fruit load is associated with a selective dampening of auxin transport in the apical region of the stem. Subsequently, an increase in auxin response in the vascular tissues of the apical stem where developing fruits are attached marks the second stage for the end of flowering transition. Similar to the vegetative and floral transition, the end of flowering transition is associated with a change in sugar signaling and metabolism in the inflorescence apex. Taken together, our results suggest that during the end of flowering transition, dominance inhibition of inflorescence shoot growth by fruit load is mediated by auxin and sugar signaling.

Dominance inhibition of inflorescence shoot growth by fruit load involves auxin and sugar signaling during the end of flowering transition.