Development of the endodermis and hypodermis ofTypha glaucaGodr. andTypha angustifoliaL. roots

摘要

The development of the endodermis and hypodermis in adventitious roots ofTypha angustifoliaL. andTypha glaucaGodr. was followed from the apical meristem to full maturity. The endodermis was typical, developing a thin Casparian band near the root tip, followed by suberin lamellae and asymmetric, secondary, lignified walls (C-type at maturity). Passage cells were present at an intermediate stage but eventually disappeared when all cells developed lamellae and secondary walls. The hypodermis was multiple (four to six layers at maturity) and began differentiating near the root tip. Here, the radial and transverse walls of the outermost layer did not dissolve in strong acid and the former were wavy in the longitudinal direction, both features characteristic of a Casparian band, but these walls were permeable to berberine. No other indication of a wall modification was seen for 3 weeks, at which time the root had become determinate and aerenchyma was beginning to form in the midcortex. Casparian bands, which were impermeable to berberine, matured in the hypodermis; thus, it proved to be an exodermis. Different forms of Casparian band were detected: one was typical and occupied the radial and transverse walls of the outermost layer, but others were novel and included tangential walls, often forming an H-shaped structure (as seen in cross section of the root). We propose calling the latter type an H-type Casparian band. It functioned as an apoplastic barrier to berberine applied either externally or internally by injection into the cortical aerenchyma. Following maturation of the Casparian band, the outer two layers of the exodermis soon produced suberin lamellae. These continued to be deposited in a centripetal pattern until they were found in all layers of the multiple exodermis. Development of the suberin wall modifications correlates with the development of the aerenchyma and may play a role in preventing gas exchange between the root and the rhizosphere. Later, all exodermal cells produced lignified, secondary walls. These were asymmetric in the outermost and innermost layers of the hypodermis (like the C-type endodermis); eventually, all layers had Casparian wall materials. Thus, the mature hypodermis consisted of two to six layers of exodermis, except at the tips of determinate roots where the exodermis was uniseriate with typical Casparian bands.Key words: cattail, endodermis, exodermis, hypodermis, roots,Typha.