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花期早晚对于植物能否在适宜的环境条件下完成生命周期具有重要意义。对于大部分观赏植物而言，花是最主要的经济器官，花的开放时间直接决定了其观赏价值和市场效益。随着花期调控技术的不断进步，观赏植物能突破自然花期限制在消费旺季集中开放，显著提升了市场竞争力和经济价值。例如通过低温诱导等技术可把蝴蝶兰Phalaenopsis开花时间从自然花期的3—5月调至春节前，使蝴蝶兰成为“年宵花之王”[1]。此外，菊花Chrysanthemum × morifolium自然花期集中在10—11月，利用其短日照诱导开花的特性，在8月初开始遮光处理，可将开花时间提早至国庆，而通过长日照补光则能延至元旦上市[2−4]。植物成花不仅受环境因子的调控，还与内部生理状态密切相关，同时涉及复杂的基因调控网络[5−8]。近年来，观赏植物栽培技术和成花机制研究不断深入，为花期精准调控提供了理论依据。本研究结合国内外最新研究成果，系统综述了观赏植物花芽分化进程、不同成花途径的分子机制及花期调控技术，以期为实现观赏植物精准花期控制及商品化生产提供理论依据。
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花芽分化是直接决定观赏植物开花时间的关键过程[9]，目前其形态变化研究已取得一定进展[5]。在此过程中植物体顶端分生组织的形态发生显著变化，包括生长点隆起或凹陷、细胞分裂活性增强及芽鳞结构改变等，生长锥的形态转变及花芽内部器官原基的发育状态是判断花芽分化阶段的核心标准[10−11]。但由于花器官结构和发育顺序的差异，不同物种花芽分化阶段划分存在差异。通常可将花芽分化过程划分为未分化期、花原基分化期、花被分化期以及雌雄蕊分化期[12−14]。而兰科Orchidaceae和菊科Asteraceae等具有花序结构的植物，其花芽分化过程较为复杂，还包含花序分化阶段[15−19]。菊科植物的典型特征是头状花序，其花芽分化始于花原基分生组织的发育，分化初始生长锥伸长变大，表面细胞层数明显增加，苞片原基形成后，按向心顺序形成小花原基，并进一步分化为舌状花和管状花[17−18]。兰科植物也因其独特的花部器官结构，分化过程需再经历合蕊柱及花粉块分化2个特殊时期[19]。以蝴蝶兰为例，花芽分化自顶端分生组织启动开始进入花序原基分化期，生长锥膨大隆起；随后从花序分生组织的侧翼分化出圆球状凸起进入花原基分化期；最终形成萼片、花瓣、蕊柱和花粉块后，完成分化[10]。而月季Rosa chinensis等单生花植物的花芽分化则更为简单，其顶端分生组织直接转化为单一花原基，无需构建花序结构[20−22]。
此外，不同观赏植物花芽分化的起始时间也存在明显差异(表1)，桂花Osmanthus fragrans、荷花Nelumbo nucifera、白玉兰Yulania denudata和杜鹃Rhododendron simsii花芽分化开始时间集中在上半年；芍药Paeonia lactiflora、寒兰Cymbidium kanran、墨兰Cymbidium sinense和大百合Cardiocrinum giganteum则集中在下半年。其中，部分植物花芽分化持续时间长达3个月以上，如桂花、牡丹Paeonia suffruticosa、芍药、春兰、胼胝兜兰Paphiopedilum callosum、大百合、山茶花Camellia japonica；部分则短至1个月左右，如郁金香属Tulipa植物。综上所述，不同观赏植物的花芽分化时间和动态发育过程都存在较大差异，直接塑造了其开花时间的多样性。因此，探究花芽分化过程对于实现花期精准调控至关重要。
表 1 观赏植物的花芽分化过程及开花时间
Table 1. Process of flower bud differentiation and the flowering time in ornamental plants
观赏植物 花芽分化开始时间 花芽分化持续时间/d 自然花期 参考文献 桂花‘厚瓣金桂’Osmanthus fragrans‘Houban Jingui’ 4月 135 9—10月 [23] 荷花Nelumbo nucifera 4—6月 3~5 6—8月 [21] 牡丹Paeonia suffruticosa 7月 90~120 4—5月 [24] 芍药Paeonia lactiflora 11月 140 4—5月 [25] 郁金香属Tulipa 7月 30~45 3—5月 [22, 26] 寒兰Cymbidium kanran 8—10月 45~60 10—12月 [27] 墨兰Cymbidium sinense 9—10月 120~150 2—3月 [28] 春兰Cymbidium goeringii 5月 120 2—3月 [29] 胼胝兜兰Paphiopedilum callosum 3月 360 4—5月 [19] 夏菊‘优香’Chrysanthemum × morifolium‘Yuuka’ 4—5月 25~44 6—9月 [16] 秋菊‘黄金球’Chrysanthemum × morifolium‘Golden Ball’ 8—9月 30 11月 [18] 大百合Cardiocrinum giganteum 11月 150 6—7月 [12] 八仙花‘经典红’Hydrangea macrophylla‘Classic Red’ 9—10月 120 6—7月 [30] 白玉兰Yulania denudata 5月 41 2—3月 [31−32] 杜鹃Rhododendron simsii 6月 120 3—5月 [33] 山茶花‘烈香’Camellia japonica‘High Fragrance’ 5月 120 12月至翌年3月 [14] -
从生物学本质来看，植物开花是先在营养生长阶段完成“幼年期—成年期”的发育转变，再通过整合外部环境刺激与内源发育信号，启动花芽分化并最终完成开花，这一过程依赖精密复杂的调控网络实现[34−36]。目前已知的花期调控路径主要包括光周期、赤霉素、环境温度、春化、自主和年龄等6条分子途径，这些途径在模式植物中已形成清晰的网络，但对于观赏植物花期调控的研究较少，且呈现显著的物种特异性和环境响应复杂性。
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光是植物光合作用的能量来源，同时也影响其成花。在光周期途径中，植物叶片会响应内源和外源信号，决定何时从营养生长向生殖生长转变[35]。其中，CONSTANS (CO)是光周期途径中的关键基因，它通过调控成花素Flowering Locus T (FT)和开花整合子Suppressor of Overexpression of CONSTANS 1 (SOC1)，进而激活APETALA1 (AP1)和LEAFY (LFY)的表达，最终形成花分生组织，实现营养生长向生殖生长的转变[36]。
根据自然花期，菊花可分为夏菊、秋菊和寒菊等，秋菊通常需要严格的短日照诱导才能完成开花，而夏菊一般对光周期不敏感[3]。在夏菊‘优香’Chrysanthemum × morifolium ‘Yuuka’中，FT-like基因FLOWERING LOCUS T-like 1(CmFTL1)被证实是在长日照下促进开花的成花素，RADICAL-INDUCED CELL DEATH 1 (CmRCD1)通过负调控与CO相同家族的B-BOX DOMAIN PROTEIN 8 (CmBBX8)蛋白活性，阻碍CmBBX8靶向CmFTL1启动子，从而抑制CmFTL1的表达，延迟开花[37−39]。HIGUCHI等[40]发现甘野菊Chrysanthemum seticuspe的Anti-florigenic FT (CsAFT)基因作为开花抑制子参与光周期路径。GIGANTEA (CsGI)在日中性条件下上调CsAFT的表达[41]，而CsAFT与FD-like1(CsFDL1)相互作用，直接拮抗CsFTL3的成花诱导活性，从而延迟花期[42−43]。NUTRITION RESPONSE AND ROOT GROWTH(CmNRRa)的表达在秋菊‘神马’Chrysanthemum × morifolium ‘Jinba’中受光周期影响，CmNRRa与14-3-3蛋白Cm14-3-3μ协同抑制CmFTL3和AP1/FRUITFULL (FUL)同源基因CmAFL1的表达，从而负调控菊花光周期开花途径[38]。菊花中CmBBX24的表达受到短日照抑制，其异源过表达的拟南芥Arabidopsis thaliana出现推迟开花的表型，且光周期路径相关基因CO、FT和SOC1均下调表达，表明CmBBX24可能通过光周期诱导途径来影响开花时间[44−45]。
此外，从文心兰Oncidium ‘Gower Ramsey’中分离得到受光周期调控的FT同源基因OnFT，该基因可以上调花分生组织特性基因AP1，并使转基因拟南芥早花[46]。春兰的CgFT基因在短日照下比长日照下表达量高，且过表达CgFT能显著上调CgLFY、CgAP1、CgFUL和SEPALLATA1 (CgSEP1)的表达，并促进开花[47−48]。在对光周期相对不敏感的月季中，光强是调控其开花的关键信号，在低光强条件下，RcPIFs在蛋白水平和转录水平都出现明显积累，并与RcCO形成复合物，干扰RcCO与RcFT启动子结合，从而抑制其表达并推迟开花；与之相反，在高光强条件下，光敏色素B (RcphyB)招募RcOST1L以促进RcPIF4降解并提早开花[49−50] (图1)。
图 1 光照介导观赏植物成花转变的分子调控网络
Figure 1. Molecular regulatory network of light-mediated floral transition in ornamental plants
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赤霉素(GAs)在植物的生长发育过程中扮演重要角色，尤其是对开花时间的调控。在拟南芥中，赤霉素能够促进生长抑制因子DELLA降解，正调控SOC1、LFY和FT的表达，促进成花转变[51]。与拟南芥不同，外源赤霉素对一次开花型月季的成花具有抑制作用，而对连续开花品种无显著影响。在春季短日照条件下，外源赤霉素可激活一次开花月季中的RoKSN基因，进而以RoKSN依赖的方式抑制RoFT、RoSOC1、RoAP1和RoLFY的表达[52−53]。但在夏季长日照环境下，施用赤霉素生物合成抑制剂多效唑并不能有效抑制RoKSN表达，也不能促进开花[54]。在菊花中，CmMYB2可通过调控赤霉素途径影响花期，过表达CmMYB2能显著提高叶片中GA的含量，上调GA合成相关基因(如CmGA20ox、CmGA3ox)并下调GA信号抑制基因CmDELLAs的表达，从而促进菊花早花[55]。在胼胝兜兰花器官分化的第5阶段，GA3能促进PcDELLAs蛋白降解以解除其对PcTCP15的抑制，同时上调PcXTH9，加速抽薹并提前开花[19]。
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环境温度途径中控制开花的关键因子是SHORT VEGETATIVE PHASE (SVP)[56]。SVP是开花抑制因子，能够与FT启动子结合进而抑制FT基因的表达[57]。在台湾蝴蝶兰Phalaenopsis aphrodite中，PaFT1在诱导开花的低温处理期间会特异性上调，且可抑制由SVP过表达引起的开花延迟[58]。此外，PhSVP可结合PhFTs启动子区域的CArG-box元件并抑制其表达，从而影响蝴蝶兰杂交品种‘Little Gem Stripes’的成花转变[59]。在墨兰中，CsSVPs能参与低温诱导开花的调控，在过表达CsSVP3后，CsFT、CsAP1、CsSOC1等基因的表达受到显著抑制，从而在开花的早期阶段发挥作用[60]。将芍药中的PISVP基因转化拟南芥，其开花时间延迟，且FT、SOC1、LFY、AP1等开花促进因子的表达量显著下调[61]。此外，CmSVP也可能通过环境温度途径抑制菊花成花[62−63]。野菊Chrysanthemum indicum的Nuclear Factor YC (CiNF-YC)家族中的CiNF-YC3与环境温度途径密切相关，过表达CiNF-YC3的拟南芥植株中，SVP表达量上调，开花明显延迟[64]。转录因子OfWRKY17能响应高温环境并促进OfC3H49的表达，且OfWRKY17-OfC3H49模块能够通过抑制OfSOC1B的表达成为桂花开花的负调节因子[65]。梅花Prunus mume中PmRGL2/PmFRL3蛋白复合体能够响应低温积累，并通过上调PmSVP和PmSVP-like抑制其开花[6] (图2) 。
图 2 温度介导观赏植物成花转变的分子调控网络
Figure 2. Molecular regulatory network of temperature-mediated floral transition in ornamental plants
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许多植物需要长时间的低温积累(即春化)才能开花。参与春化作用的关键基因是FLOWERING LOCUS C (FLC)和FRIGIDA (FRI)[66]。在春化作用发生之前，FRI通过上调表达FLC来推迟植物成花[67]。而长时间的低温会逐渐抑制FLC的表达，进而释放出开花促进因子SOC1和FT，实现成花转变[8]。
在观赏植物中，有关春化作用相关的基因研究集中在菊科和兰科植物方面。在夏菊中，CmFLC-like受15 ℃低温春化诱导表达，通过直接结合CmFTL3和CmAFL1启动子的CArG-box元件，抑制其转录进而延迟花期[68−69]。在菊花‘神马’乙烯受体基因CmETR2的超表达植株中，FLC的表达量显著上调，并表现出晚花的表型，推测该基因可能通过参与春化途径调控开花[70]。金钗石斛Dendrobium nobile需要经过春化作用才能在春季开花，其关键基因AGAMOUS-like 19 (DnAGL19)的表达在低温诱导的腋芽中大幅上调，且该基因在拟南芥中的异源过表达能提早花期，证实了DnAGL19在金钗石斛春化途径中的重要功能[71]。
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与春化途径相似，自主途径也通过调控FLC基因影响AP1和LFY的表达，最终调节植物开花过程[72]。参与拟南芥自主途径的基因主要有6个：FVE、FCA、FLK、FY、FLD和FPA，这些基因都是FT的同源基因，它们之间存在复杂的相互作用，但主要作用均是抑制FLC和SVP的转录和翻译，解除其对FT和SOC1的抑制从而促进开花[73]。在菊花中，TIFY家族基因CmJAZ1-like的过表达株系出现晚花现象，且CmDRM1和CmFVE基因的表达水平显著降低，CmFLC和CmSVP的表达水平显著升高，表明CmJAZ1-like介导的开花时间调控可能依赖于自主途径[74]。此外，CmERF110通过与自主开花途径基因CmFLK互作，影响生物钟进而加速菊花的开花进程[75]。
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年龄途径是植物依靠自身年龄控制开花的一种独立调节机制，由microRNA156 (miR156)-SPLs-miR172的级联路径组成[76]。随着植物的生长，miR156表达水平下降，SPLs的积累上升，进而直接促进LFY、AP1和SOC1表达量以促进开花[77]。而miR172是SPL的靶基因，其表达受SPL调控，并进一步负调控开花抑制子AP2[78]。
月季RcSPL1蛋白的积累受上游Rc-miR156的负调控，超表达RcSPL1会显著提高下游基因RcAP1、RcFUL和RcLFY的表达量，促进月季开花[7]。此外，HUANG等[79]发现作为植物开花和昼夜节律信号之间的关键连接点，RhSPL4-RhPRR5L模块正向调控月季开花时间。miR156靶基因SPL13A通过激活年龄途径相关基因AP1和primary-MIR172，促进台湾百合Lilium formosanum茎伸长和成花转变[80]。菊花中，CmNF-YB8是调控花期的关键因子，其表达受年龄调控，可直接结合菊花cmo-MIR156基因的启动子并调控其表达，当CmNF-YB8沉默时，cmo-MIR156表达降低、CmSPLs表达量升高，导致菊花提前完成幼年期向成年期的转变并提早开花，而异位表达cmo-MIR156可恢复CmNF-YB8沉默植株的早花表型，证实CmNF-YB8通过年龄途径中的cmo-MIR156-SPL模块调控菊花花期[81]。此外，野菊中的Cinf-yb8突变体开花时间较野生型显著提前，而CiLDL1-RNAi株系则延迟，原因是CiNF-YB8与CiLDL1通过竞争结合CiNF-YC1的组蛋白折叠结构域(HFD)分别形成功能拮抗的异源三聚体复合物，CiLDL1和CiNF-YB8对cin-MIR156ab表达具有相反的调控作用，影响该基因座上H3K4me2的水平[82]。
在复杂的成花转变调控网络中，6条调控途径并不是独立的，它们之间相互促进或制约。如RcTAF15b协同RcSPL1将年龄途径和自主途径整合，共同调节下游开花基因的表达，加速月季成花[7]；在梅花中受外源赤霉素负调控的基因PmSBP1/6与miR156f的表达模式呈负相关；在烟草中异源过表达PmSBP1/6能响应光周期调控呈现早花表型，且内源SOC1表达显著上调。上述结果表明PmSBP1/6可能同时受赤霉素和光周期途径调控[83]。此外，在不同观赏植物中，相同基因会表现出不同的功能，如过表达CmJAZ1-like基因延迟了菊花的开花，而AtJAZ1ΔJas的过表达则加速拟南芥开花[74]。
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植物能够精准感知并及时响应环境的变化，从而为开花选择最佳时机[84]。基于观赏植物成花的理论基础，对其花期调控技术展开探究，并在生产中进行了广泛应用。目前，主要通过光照调控、温度管理、植物生长调节剂调控以及不同栽培方式管理对观赏植物进行花期调控。
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光周期调控可显著影响多种观赏植物的开花时间[85]。对于短日照植物而言，适当减少光照时长能促进开花，如典型的短日照植物菊花，需要黑暗时长超过临界最小值才会开花[41]。多数兰科植物对光周期反应较弱[86]，但充足的光照有助于营养积累，进而促进开花。如随着光照时间的增加，墨兰花期延长，12 h光照处理相比于8 h光照处理能够延长墨兰花期5~10 d[28]。光质和光强对花期调控的影响亦非常重要，远红光能够在长日照条件下促进夏菊开花[87]；在强光下月季开花时间显著早于弱光[50]；30%遮光处理的低光强条件可使卡特树兰Cattleya intermedia的盛花期显著提前[88]；蝴蝶兰在低光强下花序出现时间比高光强推迟47 d[89]。
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温度对不同观赏植物花期的影响不同。许多热带兰科植物需低温诱导成花，如春兰在低于5.0 ℃，且处理时长达105~350 h时最有利于其成花[29, 90]；百合科Liliaceae植物能够通过低温春化打破种球休眠并促进花芽分化，如在4.0 ℃条件下处理百合9周能够使花期提前[91]；4.0 ℃低温处理5~7周可提前八仙花的花期，且低温时长较长时，开花天数相对延长[92]。此外，适度高温会促进兜兰花芽分化启动，如长瓣兜兰Paphiopedilum dianthum在23.3 ℃时花芽分化开始时间较16.7~17.7 ℃时提早约15 d[9]；18.0 ℃处理6周会诱导晚春时节郁金香Tulipa gesneriana子球茎从营养期向生殖期转变[93]；水仙Narcissus tazetta在30 ℃下处理80 d能够显著提高其开花率[94]。合理的温差也会影响植物开花。寒兰在昼夜温度为28 ℃/18 ℃组合下花期提前55 d，而25 ℃/15 ℃的组合延迟花期14 d[95]。
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植物生长调节剂可以调控植物体内重要的信号分子，通过调节细胞分裂、分化和器官发育过程，在花芽分化和开花进程中发挥关键作用。赤霉素可显著促进多种观赏植物的花芽分化和开花进程。对花序原基膨大期的蝴蝶兰喷施100 mg·L−1 GA3，能使蝴蝶兰开花提前5~10 d[96]；对低温处理的春兰喷施60~90 mg·L−1 GA3能进一步增强促花效果[90]；向日葵Helianthus annus在50~300 mg·L−1GA3处理下花期提前2~5 d，但持花期缩短[97]；用0.1 mmol·L−1GA3处理的菊花能在不同光周期下均提早开花[98]。值得注意的是，GA3的使用剂量需精确控制，如在八仙花中，过高或过低剂量都可能导致花序畸形或推迟开花，7.5 mg·kg−1GA3处理促花效果最好[92]。除赤霉素外，其他植物生长调节剂也参与花期调控。高浓度生长素(IAA)可抑制花梗伸长，而低浓度则促进花芽形成[99]。脱落酸(ABA)能阻碍蝴蝶兰花序形成[100]。喷施6-苄氨基嘌呤(6-BA)能刺激蝴蝶兰花芽分化，增加其花梗数和花朵数[101]，还能诱导白玉兰当年二次开花[32]。外源施加乙烯利(ETH)可使寒兰推迟开花10~20 d[95]。
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栽培管理方式(如水肥管理、修剪与摘心等)能在一定程度上影响花期。适当的氮磷钾配比能显著促进海棠‘长寿冠’Chaenomeles speciosa ‘Changshouguan’的花芽分化，当施以氮肥1.2 g·株−1、磷肥0.3 g·株−1和钾肥0.4 g·株−1时，促花效果最佳[102]。对莲瓣兰Cymbidium tortisepalum ‘Fukuyama’增施质量比为14∶20∶20的氮磷钾三元缓效复合肥，可为花器官原基形成提供充足养分，有效促进花芽分化[103]。对月季进行合理修剪可以消除其顶端抑制作用，促进侧芽萌发和花芽形成，进而促进花期[104]。此外，摘心能够打破菊花的顶端优势，使顶端的养分重新分配至侧芽，促使侧芽启动花芽分化；但二次摘心会导致整体花芽分化启动时间推迟[105]。
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花期调控作为观赏植物育种与生产的核心环节，已从传统环境调控技术向分子机制研究深度推进。借助花期调控技术及分子生物学，不仅鉴定出许多调控成花诱导与花发育时序的关键基因，也初步构建了遗传因子与环境信号互作的花期调控网络。然而相较于模式植物，观赏植物花期调控的研究仍存在显著局限：①多数观赏植物基因组信息匮乏、遗传背景复杂，且高效遗传转化体系尚未建立，导致关键基因的功能验证难以开展，阻碍了多年生花卉花期调控机制的解析；②现有研究多聚焦于单一因子对花期的影响，对于多因子的协同作用机制尚未阐明，缺乏可用于生产实践的多变量调控模型；③过度依赖温室系统的高能耗设施，成本高昂且对环境造成压力，与低碳农业的发展方向相悖。
要进一步优化花期调控的手段，未来研究需从基础机制与技术应用创新两方面进行突破，可重点关注以下几点：①挖掘特定观赏植物品种独特的花期调控通路与机制。如通过比较基因组学分析菊科中光周期敏感型品种(秋菊)与不敏感型品种(夏菊)差异，鉴定调控菊科光周期不敏感特性的关键基因，为解析观赏植物花期调控的替代途径提供线索；②强化多因子互作网络的解析，如整合环境信号、激素水平与基因表达数据，构建三者协作的花期调控模型；③突破遗传转化效率低下的瓶颈，建立高效基因编辑体系定向修饰开花关键基因，培育花期可控的新品种；④从转录水平及表观遗传角度深入解析环境因子调控观赏植物花期的分子机制，进而挖掘关键靶基因用于定向育种。这些研究可为观赏植物的分子育种与高效生产提供理论支撑，进而推动观赏花卉产业向高品质、低能耗及可持续的方向发展，最终实现“按需开花”。
Advance in flowering regulation technologies and molecular mechanisms of ornamental plants
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摘要: 开花时间作为观赏植物的重要观赏性状与经济性状，直接影响其商品价值与市场供应能力，实现花期的精准调控与周年生产已经成为观赏植物产业发展的迫切需求。因此，如何基于观赏植物的成花诱导过程及其与环境因子的关系，实现精准花期调控，对于满足市场需求具有重要意义。随着观赏植物成花机制研究和栽培技术的不断深入与进步，观赏植物花期调控的研究已取得了显著进展。本研究总结了观赏植物花芽分化的形态建成和变化规律，并系统综述了观赏植物6条成花分子路径(光周期、赤霉素、环境温度、春化、自主和年龄)及主要花期调控技术(光照、温度、植物生长调节剂、栽培方式)的最新研究进展。最后基于已有研究中存在的主要问题，提出未来的重点研究方向，以期为观赏植物精准花期调控及商品化生产提供理论支撑与实践参考。图2表1参105Abstract: As a crucial ornamental and economic trait of ornamental plants, flowering time directly affects their commercial value and market supply capacity. Achieving precise regulation of flowering time and year-round production has become an urgent demand for industrial development. Therefore, exploring how to achieve precise flowering period regulation based on the floral induction process of ornamental plants and its relationship with environmental factors holds significant importance for meeting market demands. With the continuous in-depth research on the flowering mechanism and the advancement of cultivation techniques, remarkable progress has been made in the regulation of the flowering period of ornamental plants. This paper summarized the morphological formation and developmental patterns of flower bud differentiation in ornamental plants, and systematically reviewed the latest research progress on 6 floral induction molecular pathways (photoperiod, gibberellin, ambient temperature, vernalization, autonomous, and age), and major flowering time regulation technologies (light, temperature, plant growth regulator, and cultivation method) in ornamental plants. Based on the main problems existing in the research, the key research directions for the future were proposed, to provide theoretical support and practical reference for the precise flowering period regulation and commercial production of ornamental plants. [Ch, 2 fig. 1 tab. 105 ref.]
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图 1 光照介导观赏植物成花转变的分子调控网络
Figure 1 Molecular regulatory network of light-mediated floral transition in ornamental plants
图 2 温度介导观赏植物成花转变的分子调控网络
Figure 2 Molecular regulatory network of temperature-mediated floral transition in ornamental plants
表 1 观赏植物的花芽分化过程及开花时间
Table 1. Process of flower bud differentiation and the flowering time in ornamental plants
观赏植物 花芽分化开始时间 花芽分化持续时间/d 自然花期 参考文献 桂花‘厚瓣金桂’Osmanthus fragrans‘Houban Jingui’ 4月 135 9—10月 [23] 荷花Nelumbo nucifera 4—6月 3~5 6—8月 [21] 牡丹Paeonia suffruticosa 7月 90~120 4—5月 [24] 芍药Paeonia lactiflora 11月 140 4—5月 [25] 郁金香属Tulipa 7月 30~45 3—5月 [22, 26] 寒兰Cymbidium kanran 8—10月 45~60 10—12月 [27] 墨兰Cymbidium sinense 9—10月 120~150 2—3月 [28] 春兰Cymbidium goeringii 5月 120 2—3月 [29] 胼胝兜兰Paphiopedilum callosum 3月 360 4—5月 [19] 夏菊‘优香’Chrysanthemum × morifolium‘Yuuka’ 4—5月 25~44 6—9月 [16] 秋菊‘黄金球’Chrysanthemum × morifolium‘Golden Ball’ 8—9月 30 11月 [18] 大百合Cardiocrinum giganteum 11月 150 6—7月 [12] 八仙花‘经典红’Hydrangea macrophylla‘Classic Red’ 9—10月 120 6—7月 [30] 白玉兰Yulania denudata 5月 41 2—3月 [31−32] 杜鹃Rhododendron simsii 6月 120 3—5月 [33] 山茶花‘烈香’Camellia japonica‘High Fragrance’ 5月 120 12月至翌年3月 [14] 
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