Velamen

Abstract: The velamen radicum, a spongy, usually multiple epidermis of the roots, which at maturity consists of dead cells, is frequently described as an important adaptation of epiphytic orchids. Yet, quantitative evidence for the alleged functions, e.g., efficient water and nutrient uptake, nutrient retention, reduction of water loss, mechanical protection, or the avoidance of overheating, is rare or missing. We tested the notion originally put forward by Went in 1940 that the velamen allows plants to capture and immobilize the first solutions arriving in a rainfall, which are the most heavily charged with nutrients. In a series of experiments, we examined whether all necessary functional characteristics are given for this scenario to be realistic under ecological conditions. First, we show that the velamen of a large number of orchid species takes up solutions within seconds, while evaporation from the velamen takes several hours. Charged ions are retained in the velamen probably due to positive and negative charges in the cell walls, while uncharged compounds are lost to the external medium. Finally, we demonstrate that nutrient uptake follows biphasic kinetics with a highly efficient, active transport system at low external concentrations. Thus, our results lend strong support to Went's hypothesis: the velamen fulfills an important function in nutrient uptake in the epiphytic habitat. Most of the other functions outlined above still await similar experimental scrutiny.

Abstract: Roots of representative epiphytic orchids were examined for anatomical detail, desiccation resistance and evidence of CAM activity. Those "shootless" taxa examined (Campylocentrum pachyrrhizum (Reichenb. f.) Rolfe, Harrisella porrecta Reichenb. f.) Fawc. & Rendle, and Polyradicion lindenii (Lindl.) Cogn. ex Urban) and a semi-shootless type (Kingidium taeniale (Lindl.) P. F. Hunt) bear thinner or eroded velamina and greater volumes of cortical intercellular space than do those of the leafy forms tested (Campylocentrum sellowii (Reichenb. f.) Rolfe, Encyclia tampensis (Lindl.) Small, Epidendrum radicans Pavon ex Lindl., Phalaenopsis amabilis (L.) Blume, Rangaeris amaniensis (Krzl.) Summerhayes and Vanda parishii (Reichenb. f.)). Shootless species also bear a more elaborate aeration apparatus at the velamen-cortex interface. Structurally distinct cortical cells located in this region may regulate gas exchange across the exodermis. Velamen thickness varies greatly among the ten species, as does the development of outer tangential walls of U cells in the underlying exodermis. Desiccation resistance under laboratory conditions was more closely related to root surface-to-volume ratio (S/V) than to any other measured anatomical parameter, including velamen development. Modes of carbon gain and the possible pathway for movement of fungus-borne carbon into an orchid's pool of assimilates are described, as is the possible significance of these processes to survival in forest canopy habitats. THE ORCHID PRESENCE in tropical forest canopies is unparalleled among tracheophytes. Between fifteen and twenty thousand species occur there-fully two-thirds of the family's total membership and more than all other vascular epiphytes combined (Madison, 1977). Numerous varied and complex adaptations underlie this remarkable success, but none is more basic, yet probably less appreciated, than the velamentous root. Epiphytic orchids always feature extensive specialized root systems, while their shoots sometimes consist of little more

Abstract: Summary � UV-B radiation damage in leaves is prevented by epidermal UV-screening compounds that can be modulated throughout ontogeny. In epiphytic orchids, roots need to be protected against UV-B because they photosynthesize, sometimes even replacing the leaves. How orchid roots, which are covered by a dead tissue called velamen, avoid UV-B radiation is currently unknown. � We tested for a UV-B protective function of the velamen using gene expression analyses, mass spectrometry, histochemistry, and chlorophyll fluorescence in Phalaenopsis9 hybrida roots. We also investigated its evolution using comparative phylogenetic methods. � Our data show that two paralogues of the chalcone synthase (CHS) gene family are UV-B-induced in orchid root tips, triggering the accumulation of two UV-B-absorbing flavonoids and resulting in effective protection of the photosynthetic root cortex. Phylogenetic and dating analyses imply that the two CHS lineages duplicated c. 100 million yr before the rise of epiphytic orchids. � These findings indicate an additional role for the epiphytic orchid velamen previously thought to function solely in absorbing water and nutrients. This new function, which fundamentally differs from the mechanism of UV-B avoidance in leaves, arose following an ancient duplication of CHS, and has probably contributed to the family’s expansion into the canopy during the Cenozoic.