An investigation into the role of light during desiccation of three angiosperm resurrection plants

Under water‐limiting conditions excitation energy harnessed by chlorophyll can lead to the formation of reactive oxygen species (ROS). Resurrection plants minimize their formation by preventing the opportunity for light–chlorophyll interaction but also quench them via antioxidants. Poikilochlorohyllous species such as Xerophyta humilis break down chlorophyll to avoid ROS formation. Homoiochlorophyllous types retain chlorophyll. We proposed that leaf folding during drying of Craterostigma wilmsii and Myrothamnus flabellifolius shades chlorophyll to avoid ROS (Farrant, Plant Ecology 151, 29–39, 2000). This was tested by preventing leaf folding during drying in light. As controls, plants were dried without light, and X. humilis was included. Craterostigma wilmsii did not survive drying in light if the leaves were prevented from folding, despite protection from increased anthocyanin and sucrose and elevated antioxidant enzyme activity. Membranes were damaged, electrolyte leakage was elevated and plastoglobuli (evidence of light stress) accumulated in chloroplasts. Restrained leaves of M. flabellifolius survived drying in light. Leaf folding allows less shading, but the extent of chemical protection (anthocyanin content and antioxidant activity) is considerably higher in this species compared with C. wilmsii. Chemical protection appears to be light regulated in M. flabellifolius but not in C. wilmsii. Drying in the dark resulted in loss of viability in the homoiochlorophyllous but not the poikilochlorophyllous species. It is hypothesized that some of the genes required for protection are light regulated in the former.     

Protection mechanisms against excess light in the resurrection plants Craterostigma wilmsii and Xerophyta viscosa

Mechanisms of avoidance and protection against light damage were studied in the resurrection plants Craterostigma wilmsii and Xerophyta viscosa.In C. wilmsii, a combination of both physical and chemical changes appeared to afford protection against free radical damage. During dehydration leaves curled inwards, and the abaxial surface became exposed to light. The tissue became purple/brown in colour, this coinciding with a three-fold increase in anthocyanin content and a 30% decline in chlorophyll content. Thus light-chlorophyll interactions are progressively reduced as chlorophyll became masked by anthocyanins in abaxial layers and shaded in the adaxial layers. Ascorbate peroxidase (AP) activity increased during this process but declined when the leaf was desiccated (5% RWC). During rehydration leaves uncurled and the potential for normal light-chlorophyll interaction was possible before full hydration had occurred. Superoxide dismutase (SOD) and glutathione reductase (GR) activities increased markedly during this stage, possibly affording free radical protection until full hydration and metabolic recovery had occurred.In contrast, the leaves of X. viscosa did not curl, but light-chlorophyll interactions were minimised by the loss of chlorophyll and dismantling of thylakoid membranes. During dehydration, free radical protection was afforded by a four-fold increase in anthocyanin content and increased activities of AP, GR and SOD. These declined during rehydration. It is suggested that potential free radical damage may be avoided by the persistence of anthocyanins during the period of thylakoid membrane re-assembly and full chlorophyll restitution which only occurred once the leaves were fully rehydrated.     

Ecophysiological responses of homoiochlorophyllous and poikilochlorophyllous desiccation tolerant plants: a comparison and an ecological perspective

There is an apparently stark contrast in ecophysiological adaptation between the poikilochlorophyllous desiccation-tolerant (PDT) angiosperm Xerophyta scabrida and homoichlorophyllous desiccation-tolerant (HDT) lichens and bryophytes. We summarise measurements on Xerophyta and on the temperate dry-grassland lichen Cladonia convoluta and the moss Tortula ruralis through a cycle of desiccation and rehydration. Considered in a broad ecological and evolutionary context, desiccation tolerance in general can be seen as evading some of the usual problems of drought stress, and these plants as particular instances drawn from an essentially continuous spectrum of adaptive possibilities – related on the one hand to the physical scale of the plants, and on the other to the time-scale of wetting and drying episodes.