Study on the flowering biological characteristics of three Rhododendron species
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摘要:
为提高杜鹃（Rhododendron）杂交育种效率，采用实地观测法并结合显微亚显微观测对映山红（R. simsii Planch.）、溪畔杜鹃（R. rivulare Hand.-Mazz.）和毛棉杜鹃（R. moulmainense Hook.）的开放动态、花部特征、不同开放时期花粉活力及柱头可授性等开花生物学特性进行研究。结果显示：（1）3种杜鹃花期存在不遇的情况，映山红花期为3月中旬至4月初，20~25 d，单花花期11~15 d；溪畔杜鹃花期为4月中旬至5月中旬，约30 d，单花花期12~15 d；毛棉杜鹃花期为4月中旬至5月初，15~20 d，单花花期8~13 d。（2）3种杜鹃雄蕊形态特征相似，颜色存在较大差异；花粉均为复合型四合体，表面具粘丝，可将花粉连接成串丝状；映山红花粉活力最强时期为花冠平展期（82.77%），溪畔杜鹃为喧蕾期（55.66%），毛棉杜鹃则在初开期花粉活力达到最高（81.86%）。（3）3种杜鹃花柱形态特征相似，颜色存在较大差异；柱头表面乳突细胞随开花进程均呈先皱缩而后逐渐平整饱满到最后干瘪、解体的变化；柱头可授性均呈先增强后减弱的趋势，溪畔杜鹃在初开期就具有较强可授性，映山红和毛棉杜鹃均在花冠平展期有强可授性；均可根据柱头形态特征判断可授性强弱。（4）3种杜鹃均为雄蕊先熟，雌蕊后熟。
Abstract:To enhance the efficiency of cross-breeding in Rhododendron, the flowering biological characteristics of three Rhododendron species were studied through field observation and microscopic/submicroscopic analyses. The study focused on flowering dynamics, flower traits, pollen viability, and stigma receptivity at different developmental stages. Results showed that: (1) The three Rhododendron species exhibited asynchronous flowering periods, with R. simsii flowering from mid-March to early April (approximately 20 to 25 d) and single flowers lasting 11–15 d, R. rivulare flowering from mid-April to mid-May (approximately 30 d) and single flowers lasting 12–15 d, and R. moulmainense flowering from mid-April to early May (approximately 15–20 d) and single flowers lasting 8–13 d. (2) The stamen morphology of the three species was similar, although flower colors varied significantly. The pollen in all three species formed compound tetrads, and distinct morphological changes were observed at different flower opening stages. The pollen surface exhibited sticky filaments, which connected the grains into filamentous strings. The highest pollen activity in R. simsii occurred during the petal flat extension period, reaching 82.77%, while R. rivulare achieved 55.66% viability in the bud stage and R. moulmainense reached 81.86% viability in the ready-to-open period. (3) The style morphology was similar across the three species, but notable color differences were present. Throughout development, the anthers and stigmas of all three species displayed wrinkled surface cells, which gradually became plumper, flatter, and eventually disintegrated. Stigma receptivity was initially strong but decreased over time. The stigma of R. rivulare exhibited strong receptivity at the beginning of the ready-to-open stage, while the stigmas of R. simsii and R. moulmainense exhibited strong receptivity during the petal flat extension period. Receptivity could be assessed based on morphological changes. (4) In all three species, the stamens matured before the pistils, with the latter ripening later.
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Keywords:
- Rhododendron /
- Flowering phenology /
- Stigma receptivity /
- Pollen viability
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图 1 3种杜鹃开花状态及花部结构
A列为映山红；B列为溪畔杜鹃；C列为毛棉杜鹃。
Figure 1. Flowering state and floral structure of three Rhododendron species
Column A is Rhododendron simsii; column B is Rhododendron rivulare; column C is Rhododendron moulmainense.
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图 2 3种杜鹃不同开放时期
A行为映山红；B行为溪畔杜鹃；C行为毛棉杜鹃。A：松蕾期；B：露色期；C：喧蕾期；D：初开期；E：花冠平展期；F：花冠凋落期。下同。标尺为1 cm。
Figure 2. Three Rhododendron species at different opening stages
Line A is Rhododendron simsii; line B is Rhododendron rivulare; line C is Rhododendron moulmainense. A: Pine bud stage; B: Dewness stage; C: Bud stage; D: Ready-to-open period; E: Petal flat extension period; F: Corolla falling stage. Same below. Bar=1 cm.
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图 3 3种杜鹃不同开放时期花部特征变化趋势
Figure 3. Trends in flowering characteristics of three Rhododendron species at different opening stages
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图 4 3种杜鹃不同开放时期雄蕊变化趋势
Figure 4. Trends in stamens of three Rhododendron species at different opening stages
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图 5 映山红单花不同开放时期花药及花粉的亚显微状态
A-1~A-6：松蕾期；B-1~B-6：露色期；C-1~C-6：喧蕾期；D-1~D-6：初开期；E-1~E-6：花冠平展期；F-1~F-6：花冠凋落期。A-1和A-2、B-1和B-2、C-1和C-2、D-1和D-2、E-1和E-2、F-1和F-2均为花药；A-3和A-4、B-3和B-4、C-3和C-4、D-3和D-4、E-3和E-4、F-3和F-4均为花药表皮细胞；A-5和A-6、B-5和B-6、C-5和C-6、D-5和D-6、E-5和E-6、F-5和F-6均为花粉群体；A-7、B-7、C-7、D-7、E-7、F-7均为花粉。下同。
Figure 5. Submicroscopic states of anthers and pollen of Rhododendron simsii at different opening stages
A-1 to A-6: Pine bud stage; B-1 to B-6: Dewness stage; C-1 to C-6: Bud stage; D-1 to D-6: Ready-to-open stage; E-1 to E-6: Petal flat extension stage; F-1 to F-6: Corolla falling stage. A-1 and A-2, B-1 and B-2, C-1 and C-2, D-1 and D-2, E-1 and E-2, F-1 and F-2: Anther; A-3 and A-4, B-3 and B-4, C-3 and C-4, D-3 and D-4, E-3 and E-4, F-3 and F-4: Anther epidermal cell; A-5 and A-6, B-5 and B-6, C-5 and C-6, D-5 and D-6, E-5 and E-6, F-5 and F-6: Pollen group; A-7, B-7, C-7, D-7, E-7, F-7: Pollen. Same below.
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图 6 溪畔杜鹃单花不同开放时期花药及花粉的亚显微状态
Figure 6. Submicroscopic states of anthers and pollen of Rhododendron rivulare at different opening stages
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图 7 毛棉杜鹃单花不同开放时期花药及花粉的亚显微状态
Figure 7. Submicroscopic states of anthers and pollen of Rhododendron monlmainense at different opening stages
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图 8 3种杜鹃不同开放时期花粉活力
不同大写字母表示不同杜鹃同一开放时期差异显著（P<0.05）；不同小写字母表示同种杜鹃不同开放时期差异显著（P<0.05）。
Figure 8. Pollen activity of three Rhododendron species at different opening stages
Different capital letters indicate difference in the same opening stage of different varieties (P<0.05); Different lowercase letters indicate significant difference in different opening stages of the same variety (P<0.05).
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图 9 3种杜鹃不同开放阶段花柱及柱头直径变化
Figure 9. Diameter of flower column and stigma changes in three Rhododendron species at different opening stages
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图 10 映山红单花不同开放阶段柱头与花柱亚显微状态
A-1~A-7：松蕾期；B-1~B-7：露色期；C-1~C-7：喧蕾期；D-1~D-7：初开期；E-1~E-7：花冠平展期；F-1~F-7：花冠凋落期。A-1和A-2、B-1和B-2、C-1和C-2、D-1和D-2、E-1和E-2、F-1和F-2均为柱头；A-3和A-4、B-3和B-4、C-3和C-4、D-3和D-4、E-3和E-4、F-3和F-4均为花柱表皮细胞；A-5、B-5、C-5、D-5、E-5、F-5均为柱头表面；A-6和A-7、B-6和B-7、C-6和C-7、D-6和D-7、E-6和E-7、F-6和F-7均为柱头乳突细胞；下同。
Figure 10. Submicroscopic state of style and stigma of Rhododendron simsii at different opening stages
A-1 to A-7: Pine bud stage; B-1 to B-7: Dewness stage; C-1 to C-7: Bud stage; D-1 to D-7: Ready-to-open stage; E-1 to E-7: Petal flat extension stage; F-1 to F-7: Corolla falling stage. A-1 and A-2, B-1 and B-2, C-1 and C-2, D-1 and D-2, E-1 and E-2, F-1 and F-2: Stigma; A-3 and A-4, B-3 and B-4, C-3 and C-4, D-3 and D-4, E-3 and E-4, F-3 and F-4: Style epidermal cell; A-5, B-5, C-5, D-5, E-5, F-5: Stigma surface; A-6 and A-7, B-6 and B-7, C-6 and C-7, D-6 and D-7, E-6 and E-7, F-6 and F-7: Stigma papilla cell. Same below.
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图 11 溪畔杜鹃单花不同开放阶段柱头与花柱亚显微状态
Figure 11. Submicroscopic state of style and stigma of Rhododendron rivulare at different opening stages
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图 12 毛棉杜鹃单花不同开放阶段柱头与花柱亚显微状态
Figure 12. Submicroscopic state of style and stigma of Rhododendron moulmainense at different opening stages
表 1 不同开放阶段柱头可授性
Table 1 Stigma receptivity at different opening stages
时期
Stage映山红
Rhododendron simsii溪畔杜鹃
Rhododendron rivulare毛棉杜鹃
Rhododendron moulmainense柱头形态特征
Morphological characteristics of stigma柱头可授性
Stigma receptivity柱头形态特征
Morphological characteristics of stigma柱头可授性
Stigma receptivity柱头形态特征
Morphological characteristics
of stigma柱头可授性
Stigma receptivity松蕾期 纺锤状，绿色；乳突细胞未发育完全 +/– 呈纺锤状，深绿色；乳突细胞干瘪皱缩 +/– 纺锤状，嫩绿色；乳突细胞干瘪 +/– 露色期 绿色，部分开始向上凸起；乳突细胞中间部分开始向上膨大 + 颜色略变浅，柱头成半椭圆形，开始分泌粘液；乳突细胞向上凸起 + 深绿色，柱头部分向上凸起；乳突细胞也向上凸起并伸长成各种不规则形状 + 喧蕾期 稍带红色，已有少量粘液分泌；乳突细胞成斑块状向上凸起 ++ 淡绿色，向内收缩；乳突细胞继续膨大 ++ 有少量粘液附着，柱头浅绿色；乳突细胞继续伸长膨大 ++ 初开期 红色，表面粘液分泌增多，形成一层蛋白质薄膜；乳突细胞继续向上膨大 ++ 部分呈半球状，黄绿色，分泌大量粘液；乳突细胞向上伸长 +++ 继续向上凸起，分泌大量粘液，黄绿色；乳突细胞继续向上凸起 ++ 花冠平展期 深红色，大量粘液分布；乳突细胞发育完全 +++ 黄绿色，每一裂表面逐渐平宽，有明显粘液沉淀；乳突细胞呈椭圆状 +++ 表面有明显粘液沉积，柱头米黄色，边缘逐渐干硬；乳突细胞发育完全 +++ 花冠凋落期 表面开始变干，基本无粘液或有少量粘液分布；乳突细胞解体 + 萎蔫，边缘干枯，黄白色，粘液减少；乳突细胞萎缩干瘪 + 萎蔫变褐，表面干枯；乳突细胞解体 + 注：“–”表示柱头不具可授性；“+/–”指部分柱头具可授性；“+”表示柱头具可授性；“++ ”表示柱头可授性较强；“+++”表示可授性最强。 Notes: “–” means stigma has no receptivity; “+/–” means part of stigma has receptivity; “+” means stigma has receptivity; “++ ” means stigma has strong receptivity; “+++” means stigma has the highest receptivity. 
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