Phenology and Morphology of Flower Bud Differentiation of Dexing Raspberry Bushes
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摘要:
目的 了解德兴覆盆子(掌叶覆盆子)花芽分化的形态过程及其开花物候期,为调控其花芽分化,提高德兴覆盆子的果实产量和品质提供理论依据。 方法 采用野外定点定期观测和石蜡切片法分别对德兴覆盆子的物候期和花芽分化过程的形态学进行研究。 结果 德兴覆盆子的物候期分为萌动期(芽萌动、芽膨大、芽开放)、展叶期、开花期(现蕾、始花、盛花、末花)、果实发育期(坐果、果实膨大、果实成熟、种子成熟)、结果老枝枯死期、新枝花芽形态分化期、落叶期(始落、末落)、休眠期。其中花芽形态分化期为8月初至11月中下旬,8~9月份为花芽形态分化前期,10月初花芽分化出萼片原基,10月中旬分化出花瓣原基,11月初分化出雄蕊原基,11月中旬分化出雌蕊原基,且芽体生长逐渐缓慢,进入休眠期。 结论 与多数悬钩子属植物不同,德兴覆盆子花芽寒冬休眠之前已完成形态分化,其花芽分化过程外部形态特征与内部发育存在对应关系,利用其对应关系可简单快速判断花芽分化时期,预估花期。 Abstract:Objective Phenology and morphology of flower bud differentiation were studied for information on flowering regulation as well as berry yield and quality improvements on Dexing raspberry bushes (Rubus chingii). Method The plant phenology was observed in the field, and the morphogenesis of the flower bud differentiation examined under a microscope using paraffin section. Result The entire plant growth was classified into sprouting, leaf extending, blooming, fruit developing, two-year-old branch withering, flower bud differentiating, leaf falling and dormant phases. The flower bud differentiation of the bushes occurred largely from August to mid-to-late November with 5 distinct periods. The buds were not differentiated in August and September, and calyx primordium began to differentiate in early October, petal primordium in mid-October, stamen primordium in early November, and pistil primordium in mid-November. The buds grew rapidly in the early stage and slowed down in the mid-to-late stage before reaching dormancy. Conclusion Unlike other Rubus spp, Dexing raspberry completed its flower bud differentiation before dormancy. The significant correlation between the external morphology and interior structure during the differentiation stage provided an indicator for easy estimating the flowering of the plants. -
Key words:
- Rubus chingii /
- phenology /
- flower bud differentiation
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图 1 德兴覆盆子花芽形态分化
注:A.形态分化前期;B~E.萼片原基分化期;F~G.花瓣原基分化,花托继续膨大;H~K.雄蕊原基分化;I~L.雌蕊原基分化期。A, G, J, K, L.标尺=200 μm;B, H.标尺=500 μm;C, D.标尺=50 μm;E, F, I.标尺=100 μm。V.生长锥;Se.萼片原基;Pe.花瓣原基;St.雄蕊原基;Pi.雌蕊原基。
Figure 1. Morphological flower bud differentiation of Dexing raspberry bushes
Note:A=morphological differentiation at early stage; B-E=sepal primordia differentiation phase; F-G=petal primordia differentiation phase, receptacle enlargement continued; H-K=stamen primordia differentiation phase; I-L=pistil primordia differentiation phase. A, G, J, K, L with scale bar=200 μm; B, H with scale bar=500 μm; C, D with scale bar=50 μm; E, F, I with scale bar=100 μm. V=vegetative cone; Se=sepal primordium; Pe=petal primordium; St=stamen primordium; Pi=pistil primordium.
图 2 混合芽发育的外部形态特点
注:A~B为形态分化前;C为萼片原基分化期;D为花瓣原基分化期;E为雄、雌蕊原基分化期;F为芽鳞打开。
Figure 2. External morphology of mixed bud differentiation of Dexing raspberry bushes
Note:A-B=pre-differentiation; C=sepal differentiation phase; D=petal differentiation phase; E=stamen and pistil differentiation phase; F=bud scale open.
表 1 德兴覆盆子物候期(2013-2014)
Table 1. Phenological phases of Dexing raspberry bushes (2013-2014)
物候期
Phenology农艺学特性
Agronomic characteristics2013 2014 持续时间
Duration time/d萌动期Sprouting 芽萌动 02-08~02-14 02-11~02-17 7 芽膨大 02-15~02-19 02-18~02-21 4~5 芽开放 02-20~02-26 02-22~02-28 7 展叶期Leaf extending phase - 02-27~03-04 03-01~03-06 6 开花期Blooming phase 现蕾 03-05~03-14 03-07~03-16 10 始花 03-15~03-19 03-17~03-24 5~7 盛花 03-20~03-26 03-25~03-29 5~7 末花 03-27~03-29 03-30~03-31 2~3 果实发育期Fruit developing phase 坐果 03-30~04-02 04-01~04-06 4~6 果实膨大期 04-03~04-26 04-07~04-28 22~24 坐果成熟 04-27~05-09 04-29~05-10 12~13 种子成熟 05-10~05-12 05-11~05-15 3~5 结果老枝枯死期Two-year-old banching withering phase - 6月至7月 6月至7月 60 新枝花芽形态分化期Flower bud differentiating phase - 8月至11月 8月至11月 60 落叶期Leaf falling phase 始落 10月中旬 10月中旬 70 末落 12月末 12月末 休眠期Dormant phase - 12月至次年2月 12月至次年2月 70 注:展叶期(全树有25%已萌动的芽展出第1片叶);现蕾期(全树有75%的植株现蕾);始花期(全树有5%的花开放);盛花期(全树有75%的花开放);末花期(全树有90%的花已开放)。
Note:Leaf extending phase (25% of sprouted buds showed a 1st leaf in plant); bud-showing phase (75% of branches on plant showed flower buds); flowering phase (5% of flower buds on plant started blooming); full blooming phase (75% of flowers bloomed on plant); end of flowering (90% of flowers on plant had bloomed).表 2 德兴覆盆子混合芽分化过程外部形态特征与内部发育对应关系
Table 2. Correlation between external morphology and interior structure of Dexing raspberry bushes during mixed bud differentiation
采芽日期
Date/(mm-dd)芽长×宽
Length×width/mm外部形态特征
External morphology内部发育时期
Interior structure08~09 1.5×0.1 芽通体呈绿色,瘦小 形态分化前期 10-01 3.4×1.1 芽鳞呈覆瓦状排列,芽鳞开始由绿色转为暗红色,肉眼看不到芽轴顶端的生长分化 萼片原基分化期 10-16 4.5×1.3 芽外变化不大,整个芽继续长大 花瓣原基分化期 11-01 6.7×1.4 芽外变化不大,整个芽继续长大 雄蕊原基分化期 11-16 7.1×1.6 此时芽生长缓慢,尚未显露角线 雌蕊原基分化期 翌年2月 9.8×1.9 芽鳞片开始分离,开始显露淡绿色的角线 花准备开放 -
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