林业科学 ›› 2020, Vol. 56 ›› Issue (4): 64-73.doi: 10.11707/j.1001-7488.20200407
所属专题: 林木育种
黄玲1,邓琳1,2,初彩华1,詹卉3,王曙光1,*
收稿日期:
2019-03-18
出版日期:
2020-04-25
发布日期:
2020-05-29
通讯作者:
王曙光
基金资助:
Ling Huang1,Lin Deng1,2,Caihua Chu1,Hui Zhan3,Shuguang Wang1,*
Received:
2019-03-18
Online:
2020-04-25
Published:
2020-05-29
Contact:
Shuguang Wang
摘要:
目的: 以元江箭竹花序与花器官为研究对象,进行形态解剖学观察研究,分析元江箭竹花序特征与开花特性及结实率低的原因,并试图通过组培技术改变其败育状况,为元江箭竹生殖生物学及育种工作提供新的理论数据。方法: 采用解剖观察的方法对花器官进行形态与解剖结构特征描述,运用石蜡切片对其雌、雄配子体的发育过程进行研究,采用TTC染色法测定花粉活力,并对野外与组培条件下花药的发育情况进行比较与分析。结果: 江箭竹花序顶生,偶具侧生,花序具总柄,由1片营养叶包裹。整个花序为单轴分枝,顶部小穗发育早于基部小穗,为有限花序,单个小穗基部小花先开放,因此属无限花序。每个小穗含4~5朵小花,小花均长0.9 cm,小花间由小穗轴相连,小穗轴顶端膨大具白色绒毛。小穗基部具颖片2枚;内、外稃对生,各1片;浆片3枚;雌蕊1枚;雄蕊3枚。浆片位于内稃与雄蕊之间,小花发育成熟,浆片吸水膨胀,花药伸出,花药为淡黄绿色或紫红色,花药具4室。未成熟花药壁从外向内依次由表皮、药室内壁、中层和绒毡层4层细胞组成。花药发育至二分体时期,绒毡层开始退化,绒毡层细胞属于腺质型。成熟花粉粒为二细胞型,花粉粒平均直径为43.92 μm,被TTC染成红色,活力为54.78%,但萌发率极低,甚至不萌发,花粉败育,其败育方式可总结为绒毡层过度发育、花药壁皱缩、花粉粒退化、空壳花粉粒4种类型。子房为2心皮1室,侧膜胎座,倒生胚珠,双珠被,柱头2裂呈羽毛状,短花柱长柱头型,雌雄同熟、异位。组培条件下,花芽可正常发育成小穗并开花,内外稃张开花药伸出小穗外。花药壁纤维层加厚明显,呈连续性,但多数花粉粒仍出现空壳现象。结论: 元江箭竹为混合花序,单轴分枝,开放型风媒花。元江箭竹属于典型的雄性不育,无法形成有效花粉是导致元江箭竹结实率低的主要原因。元江箭竹花芽在组培条件下可继续开花,花药伸出小穗外,但花粉发育仍然异常,通过组培技术无法改变其败育现象,说明元江箭竹的雄蕊败育可能并非养分不足造成的。
中图分类号:
黄玲,邓琳,初彩华,詹卉,王曙光. 元江箭竹花器官形态与解剖观察及其败育分析[J]. 林业科学, 2020, 56(4): 64-73.
Ling Huang,Lin Deng,Caihua Chu,Hui Zhan,Shuguang Wang. Morphological and Anatomical Observations of Floral Organs and Sterility Analysis of Fargesia yuanjiangensis[J]. Scientia Silvae Sinicae, 2020, 56(4): 64-73.
图1
元江箭竹花枝及花序 A.竹秆的一部分;B.花序侧生、顶生;C.营养叶包裹着发育未成熟的小穗;D.花序分枝(a.主轴;b.侧轴;c.小穗轴;d.顶端小穗发育最早;e.基部小穗发育最晚;f.小穗基部小花先开放);E.淡黄绿色和紫红色花药。A.Culm with branches; B. Lateral and terminal inflorescence; C.The foliage leaf wrapped immature spikelet; D. Inflorescence branch (a. Spindle; b. Lateral rachilla; c.Rachilla of spikelet; d. Apical spikelet developed first; e.Basal spikelet developed latest; f. Florets of basal spikelet opened first); E. Pale yellowish-green and purplish red anther."
图2
元江箭竹小花的形态 A.颖片(箭头);B.小花;C.小穗轴;D.内、外稃(a.外稃急尖具硬毛;b.内稃具芒);E.各时期内稃背部张开情况;F.浆片位于内稃与花丝之间(箭头:浆片);G.膜质透明的浆片;H.收缩的浆片;I.雄蕊;J.雌蕊(c.花柱;d.柱头);K.褐化的雌蕊;L.内、外稃张开,花药、柱头同时伸出。A. Glume (arrow); B. Floret; C. Rachillas of spikelet; D. Lemma and palea (a. Lemma with acute tip and hard hair; b. Palea is aristate); E. The opening status of the back of the palea in each period; F. The lodicule is located between the palea and the filament (arrow: lodicule); G. Lodicule is membranous transparent; H. Deform lodicule; I. Stamens; J. Pistil (c. Style; d. Stigma); K. Browning pistil; L. Lemma and palea opened with anthers and stigmas extending out simultaneously."
图3
元江箭竹子房及花药的形态及发育 A.侧膜胎座;B, C.胚珠发育时期;D, E.孢原细胞;F.四核胚囊(a.倒生胚珠;b.双珠被; c.珠孔);G.次生造孢细胞时期(EP:表皮细胞; EN:药室内壁; ML:中层; Ta:绒毡层); H.减数分裂前期Ⅰ(Ct:药隔组织); I.减数分裂中期Ⅰ;J.减数分裂后期Ⅰ;K.减数分裂末期Ⅰ;L.二分体;M.四分体;N.单核花粉粒;O.单核靠边期;P.二核花粉粒;Q.绒毡层退化;R.间隔的纤维层。A. Parietal placentation; B, C. Ovule development stage; D, E. The ovary at archesporial cell; F. Four nuclear embryo sac (a. Anatropous ovule; b. Double integument; c. Micropyle); G. The stage of secondary sporogenous cells (EP: Epidermis; EN: Endothecium; ML: Middle layer; Ta: Tapetum); H. Prophase of meiosisⅠ(Ct: Connective tissue); I. Metaphase of meiosis I; J. Anaphase of meiosis I; K. Telophase of meiosis I; L. Dyad; M. Tetrads; N. Juvenile mononuclear pollen grains; O. The mononuclear microspores at periphery; P.Two nucleus pollen grains; Q. Tapetum degenerate; R. Interval fibrous layer."
图4
各种发育异常的花药 A.绒毡层细胞过度生长;B.花药壁皱缩,花药内物质融合成团,绒毡层及中层细胞界限不明显;C.无花粉粒;D.部分花粉粒不具细胞质和细胞核。A. Overdeveloped tapetum layers; B. Deform anthers, the substance in anther merges into a mass, and the boundary between tapetum and middle layer cells is not distinct; C. None pollen grain was observed; D. Some pollens without nuclei and cytoplasm."
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