Development of secondary branches and apical buds of Paeonia ludlowii under cultivated conditions
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摘要:
目的 确定引种栽培条件下的大花黄牡丹二次枝及其顶芽生长发育特点、引种地和原产地花芽分化进程，为大花黄牡丹的引种栽培和开发利用提供参考。 方法 以河南栾川引种栽培的成年株大花黄牡丹为对象，观测其二次枝及其顶芽的生长发育动态，并利用石蜡切片法观察大花黄牡丹在引种地（栾川和拉萨）及原产地（林芝）三地的二次枝顶芽花芽分化进程。 结果 （1）在栾川栽植的大花黄牡丹二次枝的生长期从5月上旬持续到9月中旬，其中5月上旬—6月上旬为生长高峰；但部分花枝的腋芽发育停滞，不形成二次枝。（2）栾川栽植的大花黄牡丹二次枝的生长可分为三类：第1类占比28.57%，于7月中旬形成顶芽并开始花芽分化，次年正常开花结实；第2类主要位于花枝/果枝中下部，形成顶芽但不分化，次年仅营养生长；第3类不形成顶芽，入冬后至次年春受冻干枯。（3）原产地林芝大花黄牡丹二次枝56.25%的顶芽可分化成花芽，次年开花结实，未观察到第3类二次枝。（4）在引种地和原产地，大花黄牡丹的二次枝顶芽分化过程一致，历经6个分化阶段后，最终形成含1个主花蕾、2 ~ 3个侧花蕾、腋芽原基和叶原基的复合芽，主花蕾较侧花蕾分化时间早；腋芽原基位于第3 ~ 4个叶原基基部，次年在花枝上发育形成新的二次枝。（5）引种地和原产地顶芽花芽分化起始时间和持续时间不同，引种地栾川分化起始晚，历时相对较长（88 ~ 97 d），引种地拉萨及原产地林芝花芽分化较早，历时相对较短（近70 d）。 结论 栾川引种栽培的大花黄牡丹花芽分化比例低，分化起始较晚，持续时间较长，但能够形成稳定且正常花芽分化、开花结实的二次枝，其自然环境可作为选择大花黄牡丹的引种栽培地点的参考。 Abstract:Objective This paper aims to determine the characteristics of growth and development of secondary branches and their apical buds of cultivated Paeonia ludlowii as well as the differentiation process of flower buds in the cultivated site and the original site, then provide a reference for the introduction, cultivation, development and utilization of P. ludlowii. Method Taking the adult P. ludlowii introduced and cultivated in Luanchuan, Henan Province of Central China as the object, the growth and development dynamics of the secondary branches and its apical buds were observed. And the apical bud floral differentiation process in introduction site (Luanchuan and Lhasa) and original site (Nyingchi) was observed by paraffin section method. Result (1) In Luanchuan, the secondary branches of P. ludlowii grew from early May to mid-September, and the peak period was from early May to early June. However, some axillary buds of flowering branches were stagnant and did not develop into secondary branches. (2) In Luanchuan, the secondary branches of cultivated P. ludlowii had three growth types: the type 1, which accounted for 28.57%, its apical buds formed and began to differentiate in mid-July, blossom and bear fruits normally in following year. The type 2 was mainly located in the middle and lower parts of the flowering/fruit branches, apical buds formed but did not differentiate and there was only vegetative growth in the following year. The type 3 did not form apical bud, freeze-dried during winter and following spring. (3) In original site Nyingchi, 56.25% of the apical buds on the secondary branches of P. ludlowii differentiated and blossomed and bore fruits normally in following year, the secondary branches of type 3 were not observed. (4) The differentiation process of the apical bud on the secondary branches was the same in introduction site and original site. The apical bud finally formed a compound bud with a top flower bud, 2–3 lateral flower buds, axillary bud primordiums and leaf primordiums though six differentiation stages, the top flower bud differentiated earlier than the lateral flower buds. The axillary bud primordiums located at the base of the 3rd–4th leaf primordiums, which will develop new secondary branches on the flowering branches in the following year. (5) The start time and duration of flower bud differentiation in introduction site and original site were different. In Luanchuan, P. ludlowii began to differentiate late and lasted relatively longer (88–97 d), while in Lhasa and Nyingchi, it began to differentiate earlier and lasted relatively shorter (nearly 70 d). Conclusion The flower bud differentiation ratio of cultivated P. ludlowii in Luanchuan is low, the differentiation begins late and lasts long but it could form a stable secondary branch with normal flower bud differentiation, blossom and bear fruit.The natural environment of it could be a reference for site selection to introduce and cultivate P. ludlowii. -
Key words:
- Peaonia ludlowii /
- secondary branch /
- growth dynamics /
- flower bud differentiation /
- composite bud
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图 1 栾川大花黄牡丹二次枝发育
A. 2019年9月4日取样于栾川，主花蕾萼片原基分化时，顶芽基部叶腋处可见腋芽原基；B. 2019年10月12日取样于栾川，腋芽原基分化出叶原基；C. 2020年5月6日取样于栾川，立蕾期花枝的腋芽；D&E. 2020年8月25日取样于栾川，果枝上部腋芽萌发形成二次枝，下部腋芽不萌发；ABP. 腋芽原基；ApB. 二次枝顶芽；Le. 叶原基；①. 花枝中上部腋芽；②. 花枝/果枝下部腋芽，逐渐膨大或加长，但不形成二次枝；③. ①发育成的二次枝。下同。A, sampled in Luanchuan on September 4, 2019, the axillary bud primordia in the basal leaf axils of the apical buds when the sepal primordium differentiates ; B, sampled in Luanchuan on October 12, 2019, the axillary bud primordia differentiates into leaf primordia; C, sampled in Luanchuan on May 6, 2020, the axillary buds at bud stage；D&E, sampled in Luanchuan on August 25, 2020, the axillary buds on upper part of the fruit branch develop into secondary branches, and the axillary buds on lower part are not germimated. ABP, axillary bud primordium; ApB, apical bud on secondary branch; Le, leaf primordia; ①, the secondary branch on the upper and middle part; ②, atagnant axillary bud，gradually enlarged or lengthened, failing to form a secondary branch; ③, the secondary branch developed from ①. The same below.
Figure 1. Development of secondary branch of Paeonia ludlowii in Luanchuan
图 2 栾川大花黄牡丹二次枝示意图（A）及平均生长动态（B）
BD. 二次枝基部直径；BL. 二次枝长度；FB. 果枝；Fr. 果实；④. 上部二次枝；⑤. 下部二次枝。BD, base diameter of secondary branch; BL, length of secondary branch; FB, fruit branch; Fr, frui; ④, upper secondary branch; ⑤, lower secondary branch.
Figure 2. Schematic diagram (A) and average growth dynamics (B) of secondary branches of P. ludlowii in Luanchuan
图 3 栾川大花黄牡丹二次枝顶芽平均生长动态
A. 形成顶芽的二次枝；B. 不形成顶芽的二次枝（2020年9月5日，栾川）；C. 二次枝顶芽平均生长动态。A, secondary branch with apical bud; B, secondary branch that could not form apical bud (taken on September 5, 2020，Luanchuan); C, average growth dynamics of apical buds of secondary branches.
Figure 3. Average growth dynamics of apical buds of secondary branches of P. ludlowii in Luanchuan
图 4 大花黄牡丹二次枝顶芽分化过程
A. 花芽分化初期；B. 苞片原基分化期；C. 萼片原基分化期；D. 花瓣原基分化期；E. 雄蕊原基分化期；F. 雌蕊原基分化期；G. 侧花蕾晚于主花蕾完成分化；H. 顶芽内不同位置的侧蕾分化时期不同；I. 含主花蕾、侧花蕾、叶原基、腋芽原基的复合芽。GP. 生长点；Le. 叶片原基；Br. 苞片原基；Se. 萼片原基；Pe. 花瓣原基；St. 雄蕊原基；Sta. 雄蕊：Pi. 雌蕊原基；MFB. 主花蕾；LFB. 侧花蕾；LFB1.顶芽基部向上第1个侧花蕾；LFB2. 顶芽基部向上第2个侧花蕾。A, early stage of flower bud differentiation; B, bract primordium differentiation period; C, Sepal primordium differentiation period; D, Petal primordium differentiation period; E, stamen primordium differentiation period; F, pistillode primordium differentiation period; G, the lateral flower-bud differentiated later than the main flower-bud; H, different position of lateral flower buds; I, compound buds with main flower bud, lateral flower buds, leaf primordium and axillary bud primordium. GP, growing point; Le, leaf primordium; Ax, axillary bud; Br, bract primordium; Se, sepal primordium; Pe, petal primordium; St, stamen primordium; Sta, staminode; Pi, pistil primordium; MFB, main flower bud; LFB, the lateral flower bud; LFB1, the first lateral flower bud; LFB2, the second lateral flower bud.
Figure 4. Differentiation process of the apical buds on the secondary branches of P. ludlowii
表 1 大花黄牡丹二次枝顶芽性状
Table 1. Characteristics of the apical buds on secondary branches of P. ludlowii
试验区 Test area 二次枝和未发育的腋芽在花/果枝上的着生位置
Implanting position of secondary branches and undeveloped axillary buds on flowering/fruiting branch第1类二次枝占比 Proportion of the type 1 secondary branch/% 上 Upper 中 Intermediate 下 Lower 不生成二次枝的腋芽数 Number of axillary buds that don,t develop into secondary branch 第1类二次枝数 Number of the type 1 secondary branch 第2类二次枝数 Number of the type 2 secondary branch 第3类二次枝数 Nnumber of the type 3 secondary branch 不生成二次枝的腋芽数 Number of axillary buds that don,t develop into secondary branch 第1类二次枝数 Number of the type 1 secondary branch 第2类二次枝数 Number of the type 2 secondary branch 第3类二次枝数 Nnumber of the type 3 secondary branch 不生成二次枝的腋芽数 Number of axillary buds that don,t develop into secondary branch 第1类二次枝数 Number of the type 1 secondary branch 第2类二次枝数 Number of the type 2 secondary branch 第3类二次枝数 Nnumber of the type 3 secondary branch 栾川
Luanchuan0 18 10 2 1 2 7 0 16 0 14 0 28.57 林芝
Nyingchi0 30 0 0 0 1 5 0 14 5 9 0 56.25 
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表 2 大花黄牡丹顶芽主花蕾的分化进程
Table 2. Differentiation process of the main flower buds in apical buds of P. ludlowii
试验区
Test area取样年份
Year of
sampling叶原基形成结束时间
Leaf primordium formation
ending time花芽分化起始时间
Initiation time of flower bud differentiation苞片原基
Bract primordium萼片原基
Sepal primordium花瓣原基
Petal primordium雄蕊原基
Stamen primordium雌蕊原基
Pistil primordium栾川 Luanchuan 2019 7月26日
July 268月2日
August 29月4日
September 49月23日
September 2310月12日
October 1210月31日
October 312020 7月16日
July 167月26日
July 268月14日
August 149月10日
September 1010月1日
October 110月11日
October 11拉萨 Lhasa 2019 7月17日
July 177月21日
July 218月11日
August 118月25日
August 259月8日
September 89月23日
September 23林芝 Nyingchi 2019 7月16日
July 167月23日
July 238月10日
August 108月23日
August 239月2日
September 29月20日
September 20
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