Zhang Xiaohui, Sawhney Vipen K, Davis Arthur R
College of Life Sciences, Shaanxi Normal University, Xi'an 710062, China.
Department of Biology, University of Saskatchewan, Saskatoon, Saskatchewan, S7N 5E2, Canada.
Am J Bot. 2014 Nov;101(11):1849-67. doi: 10.3732/ajb.1400368. Epub 2014 Oct 30.
Many angiosperms produce nectar that entices pollinator visits. Each floral nectary tends to embody a singular form, such as the receptacular ring arising beneath the ovary in mint flowers (Lamiaceae). Exceptionally, the annular floral nectary in Salvia farinacea possesses modified stomata plus secretory trichomes. This first study of nectary ultrastructure within the largest genus of Lamiaceae examined this unusual condition.
Nectary anatomy, histochemistry, and ultrastructure were investigated from fresh and fixed material using light microscopy and scanning electron and transmission electron microscopy.
The annular nectary encircled the ovary plus extended ventrally as a projection. Modified stomata occurred only in the projection's abaxial epidermis. Conversely, peltate trichomes with a basal cell, a stalk cell, and 4-7 head cells were interspersed among the ovary lobes and covered the projection's adaxial surface. Phloem and xylem supplied the nectary interior, where parenchyma cells had numerous mitochondria and plastids with little starch, but few dictyosomes and little endoplasmic reticulum. Nectar accumulated as a drop opposite the projection's abaxial surface, escaping through stomatal pores and probably the cuticle. However, the annular nectary's glistening trichomes secreted a Sudan-positive product largely retained below the distended cuticle, but not nectar.
This first ultrastructural study of co-occurring secretory trichomes and modified stomata on a mint nectary suggests multiple interactive functions for this atypical structure. These trichomes-possibly generating a substance informative to pollinators or as an ovarian defense against phytophagy-produced oil in an aqueous milieu, rather than contributing fluid to nectar.
许多被子植物会分泌花蜜以吸引传粉者来访。每个花蜜腺往往呈现出单一的形态,比如薄荷科(唇形科)花朵子房下方出现的花托环状蜜腺。例外的是,蓝花鼠尾草的环状花蜜腺具有特化的气孔和分泌毛状体。在唇形科最大的属内,首次对蜜腺超微结构进行的这项研究对这种特殊情况展开了调查。
利用光学显微镜、扫描电子显微镜和透射电子显微镜,对新鲜和固定材料的蜜腺解剖结构、组织化学和超微结构进行了研究。
环状蜜腺环绕着子房,并在腹侧延伸形成一个突起。特化的气孔仅出现在突起的远轴表皮。相反,具有一个基细胞、一个柄细胞和4至7个头部细胞的盾状毛状体散布在子房裂片之间,并覆盖突起的近轴表面。韧皮部和木质部为蜜腺内部提供养分,薄壁细胞中有大量线粒体和质体,淀粉含量很少,但高尔基体和内质网较少。花蜜在突起远轴表面相对的位置聚集成滴,通过气孔孔道以及可能通过角质层排出。然而,环状蜜腺上闪亮的毛状体分泌出一种苏丹阳性产物,该产物大部分保留在膨胀的角质层下方,而非花蜜。
对薄荷蜜腺上同时出现的分泌毛状体和特化气孔进行的首次超微结构研究表明,这种非典型结构具有多种相互作用的功能。这些毛状体可能产生一种对传粉者有信息价值的物质,或者作为子房抵御植食的防御机制,在水性环境中产生油脂,而不是为花蜜提供液体成分。