Cell walls of lichens in sub-order Peltigerineae have much higher activities and a greater diversity of cell wall redox enzymes compared with other lichens. Possible roles of tyrosinases include melanization, removal of toxic phenols or quinones, and production of herbivore deterrents.
Chlorophyll fluorescence was used to study the effect of a hardening treatment on aspects of desiccation tolerance in the moss Atrichum androgynum. Mosses were hardened by reducing the relative water content of apical stem segments to 0.5 for 3 days, followed by storing the material fully hydrated for 1 day. Desiccation tolerance was assessed by storing plants over silica gel for 16 h, and then monitoring the recovery of photosystem II during rehydration. Hardening decreased the efficiency of photosystem II before desiccation, particularly at saturating light intensities. During rehydration, photosystem II activity recovered faster in hardened plants. Hardening also considerably increased non-photochemical quenching during the first few hours of rehydration. Previous data had shown that photophosphorylation, but not carbon fixation, is starting at this time, suggesting that the moss needs strong protection from excess light. Increased nonphotochemical quenching was the result of greater thylacoidal-based or fast-relaxing quenching (qE), rather than reaction centre-based or slow-relaxing quenching (qI). Slow-relaxing quenching was similar in hardened and non-hardened mosses, and was only important during the early stages of rehydration when the need for photoprotection was greatest. In A. androgynum hardening apparently shifts the photosynthetic apparatus from a 'high efficiency' state to a less efficient but 'photoprotected' state.
Recovery of photosynthesis in rehydrating desiccated leaves of the poikilochlorophyllous desiccation-tolerant plant Xerophyta scabrida was investigated. Detached leaves were remoistened under 12 h light/dark cycles for 96 h. Water, chlorophyll (Chl), and protein contents, Chl fluorescence, photosynthesis–CO2 concentration response, and the amount and activity of Rubisco were measured at intervals during the rehydration period. Leaf relative water contents reached 87% in 12 h and full turgor in 96 h. Chl synthesis was slower before than after 24 h, and Chla:Chlb ratios changed from 0.13 to 2.6 in 48 h. The maximum quantum efficiency recovered faster during rehydration than the photosystem II operating efficiency and the efficiency factor, which is known to depend mainly on the use of the electron transport chain products. From 24 h to 96 h of rehydration, net carbon fixation was Rubisco limited, rather than electron transport limited. Total Rubisco activity increased during rehydration more than the Rubisco protein content. Desiccated leaves contained, in a close to functional state, more than half the amount of the Rubisco protein present in rehydrated leaves. The results suggest that in X. scabrida leaves Rubisco adopts a special, protective conformation and recovers its activity during rehydration through modifications in redox status.
This review discusses the production of reactive oxygen species (ROS) by lichens and their possible roles. All organisms produce ROS, and production is increased by many abiotic and biotic stresses. Intracellular ROS production is generally considered to be harmful, and a variety of enzymic and non-enzymic scavenging systems exist to detoxify them. However, extracellular ROS formation has been suggested to play ‘positive roles’, particularly in the response of organisms to stress. Given their high stress tolerance, it is rather surprising that studies on extracellular ROS production by lichens have just started. Surveys of a wide range of lichens have shown that constitutively high rates of extracellular superoxide production occur in the Suborder Peltigerineae, but production appears to be absent in other groups. In some members of the Peltigerineae ROS production is stimulated by desiccation and wounding. It seems probable that the enzymes that produce the superoxide are laccases, based on first the types of substrates that lichens can break down, and second the dependence of the breakdown of these substrates on pH, temperature and the presence of inhibitors. While much more work is needed, we suggest that physiological roles of extracellular ROS production will be found to include defence against pathogens, melanization, and lignin breakdown.
In our present studies, the recovery of photosynthetic activity after rehydration was demonstrated. We measured chlorophyll fluorescence, CO2 gas exchange and the pigment composition in the previously long-term air-dried cryptogamic inselberg crusts collected from two tropical areas. The cryptobiotic crusts were collected from different localities on similar ecological and climatic conditions from extreme habitats of inselbergs (outcrops). These inselbergs are characterized by a dry microclimate and are covered by scarce soil. We found that the ecophysiological responses of both cryptogamic inselberg crusts showed an extremely high degree of desiccation-tolerance due to the fast and full recovery during rehydration. The photosynthetic activity of the cryptobiotic crusts were restored and regained within 15 and 40 min, respectively, after rehydration. Photosynthetic activity of the crusts was retained at all applied light intensities when enough water was available, however the degree of the recovery was different between the crusts. Photosynthetic pigment contents were strongly and positively correlated with water content. Our results indicated that tropical desiccation-tolerant cryptogamic crusts found on inselberg rock surfaces have CO2 fixation ability in the range of cyanobacteria and lichens, suggesting that at a global scale they can assimilate CO2 in a significant amount.
In lichens, each symbiotic partner cooperates for the survival of the symbiotic association. The protection of the susceptible photosynthetic apparatus is essential for both participants. The mycobiont and photobiont contribute to the protection against the damaging effect of excess light by various mechanisms. The present study investigated the effect of seasonality and microhabitat exposure on photoprotection and photoacclimation in the photo- and the mycobiont of six lichen species with different thallus morphology in inland dune system in the Kiskunság region (Hungary) with shaded, more humid and exposed, drier dune sides. High-Performance Liquid Chromatography, spectrophotometry, chlorophyll a fluorescence kinetic technique were used, and micrometeorological data were collected. The four years data series revealed that the north-east-facing side was characterized by higher relative humidity and lower light intensities compared to the south-west-facing drier and more exposed sides. The south-west facing side was exposed to direct illumination 3–4 hours longer in winter and 1–2 hours shorter in summer than the north-east facing side of the dune, influencing the metabolism of sun and shade populations of various species. Because rapid desiccation caused short active periods of lichens during bright and drier seasons and on exposed microhabitats, the rapid, non-regulated non-photochemical quenching mechanisms in the photobiont had a significant role in protecting the photosynthetic system in the hydrated state. In dehydrated conditions, thalli were mainly defended by the solar screening metabolites produced by the mycobiont and curling during desiccation (also caused by the mycobiont). Furthermore, the efficacy of light use (higher chlorophyll and carotenoid concentration) increased because of short hydrated periods. Still, a lower level of received irradiation was appropriate for photosynthesis in dry seasons and on sun exposed habitats. In humid seasons and microhabitats, more extended active periods lead to increased photosynthesis and production of solar radiation protectant fungal metabolites, allowing a lower level of photoprotection in the form of regulated non-photochemical quenching by the photobiont. Interspecific differences were more pronounced than the intraspecific ones among seasons and microhabitat types.
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