Full sunlight acclimation mechanisms in Riccia discolor thalli: Assessment at morphological, anatomical, and biochemical levels.

Light occupies a central position in regulating development of plants. Either little or excess of light could be harmful for plants. Since bryophytes are shade loving organisms, they must adapt to function in fluctuating light regimes. Therefore, the aim of this study was to investigate acclimatory responses of Riccia discolor thalli grown under full sunlight, and were compared with shade grown thalli (control). Length, width, and fresh mass of thallus were significantly lower (by 27, 41 and 37%, respectively) but endogenous nitric oxide content (by 81%) and nitric oxide synthase like activity (by 58%) were higher in full sunlight grown thalli than shade grown thalli. Number of rhizoids was greater in shade but length and width of rhizoids were higher (by 36 and 25%, respectively) in full sunlight grown thalli. The content of carotenoids was higher (by 34%) in full sunlight grown thalli. In full sunlight grown thalli, chloroplasts exhibited avoidance movement but in shade grown thalli they exhibited accumulation movement. Photosynthetic yields were higher in shade grown thalli. Among energy fluxes, ABS/RC did not vary but DI0/RC was higher (by 12%) in full sunlight grown thalli. Reactive oxygen species and damage were greater in full sunlight grown thalli despite enhanced levels of antioxidants i.e. superoxide dismutase (by 66%) and catalase (by 34%). Overall results suggest that full sunlight acclimation in Riccia discolor thalli occurred at various levels in which endogenous NO plays a positive role.

Phototropin perceives temperature based on the lifetime of its photoactivated state.

Living organisms detect changes in temperature using thermosensory molecules. However, these molecules and/or their mechanisms for sensing temperature differ among organisms. To identify thermosensory molecules in plants, we investigated chloroplast positioning in response to temperature changes and identified a blue-light photoreceptor, phototropin, that is an essential regulator of chloroplast positioning. Based on the biochemical properties of phototropin during the cellular response to light and temperature changes, we found that phototropin perceives temperature based on the temperature-dependent lifetime of the photoactivated chromophore. Our findings indicate that phototropin perceives both blue light and temperature and uses this information to arrange the chloroplasts for optimal photosynthesis. Because the photoactivated chromophore of many photoreceptors has a temperature-dependent lifetime, a similar temperature-sensing mechanism likely exists in other organisms. Thus, photoreceptors may have the potential to function as thermoreceptors.

Land plants drive photorespiration as higher electron-sink: comparative study of post-illumination transient O2 -uptake rates from liverworts to angiosperms through ferns and gymnosperms.

In higher plants, the electron-sink capacity of photorespiration contributes to alleviation of photoinhibition by dissipating excess energy under conditions when photosynthesis is limited. We addressed the question at which point in the evolution of photosynthetic organisms photorespiration began to function as electron sink and replaced the flavodiiron proteins which catalyze the reduction of O2 at photosystem I in cyanobacteria. Algae do not have a higher activity of photorespiration when CO2 assimilation is limited, and it can therefore not act as an electron sink. Using land plants (liverworts, ferns, gymnosperms, and angiosperms) we compared photorespiration activity and estimated the electron flux driven by photorespiration to evaluate its electron-sink capacity at CO2 -compensation point. In vivo photorespiration activity was estimated by the simultaneous measurement of O2 -exchange rate and chlorophyll fluorescence yield. All C3-plants leaves showed transient O2 -uptake after actinic light illumination (post-illumination transient O2 -uptake), which reflects photorespiration activity. Post-illumination transient O2 -uptake rates increased in the order from liverworts to angiosperms through ferns and gymnosperms. Furthermore, photorespiration-dependent electron flux in photosynthetic linear electron flow was estimated from post-illumination transient O2 -uptake rate and compared with the electron flux in photosynthetic linear electron flow in order to evaluate the electron-sink capacity of photorespiration. The electron-sink capacity at the CO2 -compensation point also increased in the above order. In gymnosperms photorespiration was determined to be the main electron-sink. C3-C4 intermediate species of Flaveria plants showed photorespiration activity, which intermediate between that of C3- and C4-flaveria species. These results indicate that in the first land plants, liverworts, photorespiration started to function as electron sink. According to our hypothesis, the dramatic increase in partial pressure of O2 in the atmosphere about 0.4 billion years ago made it possible to drive photorespiration with higher activity in liverworts.

Ferns, mosses and liverworts as model systems for light-mediated chloroplast movements.

Light-induced chloroplast movement is found in most plant species, including algae and land plants. In land plants with multiple small chloroplasts, under weak light conditions, the chloroplasts move towards the light and accumulate on the periclinal cell walls to efficiently perceive light for photosynthesis (the accumulation response). Under strong light conditions, chloroplasts escape from light to avoid photodamage (the avoidance response). In most plant species, blue light induces chloroplast movement, and phototropin receptor kinases are the blue light receptors. Molecular mechanisms for photoreceptors, signal transduction and chloroplast motility systems are being studied using the model plant Arabidopsis thaliana. However, to further understand the molecular mechanisms and evolutionary history of chloroplast movement in green plants, analyses using other plant systems are required. Here, we review recent works on chloroplast movement in green algae, liverwort, mosses and ferns that provide new insights on chloroplast movement.

Sex differences and plasticity in dehydration tolerance: insight from a tropical liverwort.

BACKGROUND AND AIMS: Adaptations allowing plants to cope with drying are particularly relevant in the light of predicted climate change. Dehydration tolerance (DhT, also dehydration-tolerant) is one such adaptation enabling tissue to survive substantial drying. A great deal of work has been conducted on highly DhT species. However, bryophytes showing less intense and variable DhT are understudied, despite the potential for these species to provide an informative link between highly tolerant and sensitive species. In this study, we tested the degree to which DhT varies across populations and the sexes of a species expected to exhibit a moderate DhT phenotype. METHODS: To test predicted patterns of tolerance we assessed DhT in males and females of Marchantia inflexa from two distinct habitat types that differ in water availability. Both common garden and field-collected tissue was subjected to drying assays at multiple intensities and recovery was monitored by chlorophyll florescence. Verification studies were conducted to confirm the level of dehydration, the rate of drying and the associated changes in photosynthetic physiology. KEY RESULTS: We confirmed our expectation that M. inflexa is able to tolerate moderate dehydration. We also found that females exhibited higher DhT than males, but populations did not differ in DhT when cultured in a common garden. However, field-collected samples exhibited differences in DhT corresponding to environmental dryness, suggesting plasticity in DhT. CONCLUSIONS: By studying a less extreme DhT phenotype we gained insight into how more sensitive (yet still tolerant) organisms cope with dehydration. Additionally, the identified sex-specific variation in DhT may explain ecological patterns such as female-biased sex ratios. Furthermore, plasticity in DhT has the potential to inform management practices aimed at increasing tolerance to drought conditions.

Dynamic response of UV-absorbing compounds, quantum yield and the xanthophyll cycle to diel changes in UV-B and photosynthetic radiations in an aquatic liverwort.

We studied the diel responses of the liverwort Jungermannia exsertifolia subsp. cordifolia to radiation changes under laboratory conditions. The samples were exposed to three radiation regimes: P (only PAR), PA (PAR+UV-A), and PAB (PAR+UV-A+UV-B). The day was divided in four periods: darkness, a first low-PAR period, the high-PAR plus UV period, and a second low-PAR period. After 15 days of culture, we measured photosynthetic pigments, chlorophyll fluorescence and UV-absorbing compounds in the four periods of the day on two consecutive days. With respect to UV-absorbing compounds, we analyzed their global amount (as the bulk UV absorbance of methanolic extracts) and the concentration of seven hydroxycinnamic acid derivatives, both in the soluble (mainly vacuolar) and insoluble (cell wall-bound) fractions of the plant extracts. PAB samples increased the bulk UV absorbance of the soluble and insoluble fractions, and the concentrations of p-coumaroylmalic acid in the soluble fraction and p-coumaric acid in the cell wall. Most of these variables showed significant diel changes and responded within a few hours to radiation changes (more strongly to UV-B), increasing at the end of the period of high-PAR plus UV. F(v)/F(m), Φ(PSII), NPQ and the components of the xanthophyll cycle showed significant and quick diel changes in response to high PAR, UV-A and UV-B radiation, indicating dynamic photoinhibition and protection of PSII from excess radiation through the xanthophyll cycle. Thus, the liverwort showed a dynamic protection and acclimation capacity to the irradiance level and spectral characteristics of the radiation received.

Effects of enhanced UV-B radiation on hydroxycinnamic acid derivatives extracted from different cell compartments in the aquatic liverwort Jungermannia exsertifolia subsp. cordifolia.

We examined the responses of ultraviolet-absorbing compounds (UVAC) to enhanced UV-B radiation in the aquatic liverwort Jungermannia exsertifolia subsp. cordifolia for 31 days under laboratory conditions. Samples were exposed to three radiation regimes: P (only photosynthetic radiation), PA (photosynthetic + UV-A radiation) and PAB (photosynthetic + UV-A + UV-B radiation). We measured both the bulk UV absorbance of the methanolic extracts and the levels of individual UVAC. In both cases, the methanol-soluble and the methanol-insoluble, alkali-extractable cell wall-bound fractions were analyzed. The bulk UV absorbance of the soluble fraction was higher than that of the cell wall-bound fraction. The bulk UV absorbances of both fractions increased under enhanced UV-B (PAB regime). Five different hydroxycinnamic acid (HCA) derivatives were found in the soluble fraction and two additional ones in the cell wall-bound fraction, among which only p-coumaroylmalic acid in the soluble fraction and p-coumaric acid in the cell wall-bound fraction increased under enhanced UV-B. The maximum quantum yield of PSII (F(v) /F(m)) decreased and DNA damage (amount of thymine dimers) strongly increased under enhanced UV-B, showing UV-B-induced damage. We conclude that methanol-soluble and cell wall-bound fractions of the liverwort studied have different UVAC, and each individual compound may respond in a different way to UV-B radiation. Thus, the analysis of individual UVAC in both the methanol-soluble and cell wall-bound fractions is advisable to better evaluate the protection mechanisms of liverworts against UV-B radiation. In particular, p-coumaric acid and p-coumaroylmalic acid seem to be especially UV-B responsive and merit further investigation.

Retrospective bioindication of stratospheric ozone and ultraviolet radiation using hydroxycinnamic acid derivatives of herbarium samples of an aquatic liverwort.

We analyzed bulk UV absorbance of methanolic extracts and levels of five UV-absorbing compounds (hydroxycinnamic acid derivatives) in 135 herbarium samples of the liverwort Jungermannia exsertifolia subsp. cordifolia from northern Europe. Samples had been collected in 1850-2006 (96% in June-August). Both UV absorbance and compound levels were correlated positively with collection year. p-Coumaroylmalic acid (C1) was the only compound showing a significant (and negative) correlation with stratospheric ozone and UV irradiance in the period that real data of these variables existed. Stratospheric ozone reconstruction (1850-2006) based on C1 showed higher values in June than in July and August, which coincides with the normal monthly variation of ozone. Combining all the data, there was no long-term temporal trend from 1850 to 2006. Reconstructed UV showed higher values in June-July than in August, but again no temporal trend was detected in 1918-2006 using the joint data. This agrees with previous UV reconstructions.

Seasonal variations in UV-absorbing compounds and physiological characteristics in the aquatic liverwort Jungermannia exsertifolia subsp. cordifolia over a 3-year period.

Temporal physiological variations in relation to environmental factors, in particular to ultraviolet (UV) radiation, have been studied in bryophytes from circumpolar latitudes, but not in mid-latitudes with longer growing seasons. In addition, seasonal and interannual changes in individual UV-absorbing compounds (UVAC) have not been previously studied in bryophytes. To fill these gaps, samples of the aquatic liverwort Jungermannia exsertifolia subsp. cordifolia were collected on a monthly basis during 3 years from a mountain stream in northern Spain. Sclerophylly index, chlorophyll fluorescence, DNA damage, the bulk UV absorbance of methanolic extracts and the concentration of five UVAC (hydroxycinnamic acid derivatives) were measured. Interannual changes were little marked, probably because the 3 years studied were environmentally similar. In summer-autumn, with respect to seasonal variations, newly grown tender young shoots with high F(v)/F(m) accumulated higher amounts of several hydroxycinnamic acid derivatives than in winter-spring. DNA damage was not detected in any of the samples analyzed. p-Coumaroylmalic acid was the compound best associated with radiation changes, and the best model explaining UV-B took into consideration the concentration of this compound and the ozone level. The specific effects of UV radiation could not be separated from the effects caused by other environmental factors, such as global radiation or temperature, because all these variables were correlated. However, indirect evidence strongly suggests that seasonal changes in bulk UV absorbance and p-coumaroylmalic acid are mainly driven by UV radiation. This compound may be a promising physiological variable to be used for UV bioindication.

Genistein and changes of resting and action potentials in Conocephalum conicum.

Genistein (4',5,7-trixydroxyflavone) is a member of the family of plant flavonoids that widely occurs in crop plants. It is involved in a wide spectrum of pharmacological activities, and is suggested to have anti-cancer dietary properties. Cell membranes are one of the targets of anti-cancer drugs. In the present study, we used the liverwort Conocephalum conicum as a model plant in an electrophysiological study. Intracellular microelectrode measurements were carried out to examine the effects of genistein alone and in combination with verapamil on resting and action potentials. The application of isoflavone genistein resulted in a statistically significant elevation in action potential amplitudes. An increase of 13-62% compared with the control was noted. An increase was also found in the membrane resting potentials in genistein-treated plants. Verapamil, the known calcium channel inhibitor, caused a gradual decline of AP amplitudes, whereas preincubation of Conocephalum thalli with genistein prevented inhibition of APs by verapamil. It is concluded that genistein strongly affects the membranes, and the effect of genistein in canceling the activity of verapamil is discussed.

Hydroxycinnamic acid derivatives in an aquatic liverwort as possible bioindicators of enhanced UV radiation.

We examined, under laboratory conditions, the physiological responses of the aquatic liverwort Jungermannia exsertifolia subsp. cordifolia to artificially enhanced ultraviolet (UV) radiation for 82 days, especially considering the responses of five hydroxycinnamic acid derivatives. This species lives in mountain streams, where it is exposed to low temperatures and high UV levels, and this combination is believed to increase the adverse effects of UV. Enhanced UV radiation hardly caused any change in several physiological variables indicative of vitality, such as Fv/Fm and chlorophylls/phaeopigments ratio (OD430/OD410). Thus, this liverwort seemed to be tolerant to UV radiation, probably due to the accumulation of three UV-absorbing hydroxycinnamic acid derivatives: p-coumaroylmalic acid, 5''-(7'',8''-dihydroxycoumaroyl)-2-caffeoylmalic acid, and 5''-(7'',8''-dihydroxy-7-O-beta-glucosyl-coumaroyl)-2-caffeoylmalic acid. These compounds might serve as bioindicators of enhanced UV radiation.

Effects of cadmium and enhanced UV radiation on the physiology and the concentration of UV-absorbing compounds of the aquatic liverwort Jungermannia exsertifolia subsp. cordifolia.

The aquatic liverwort Jungermannia exsertifolia subsp. cordifolia was cultivated for 15 d under controlled conditions to study the single and combined effects of cadmium and enhanced ultraviolet (UV) radiation. Both cadmium and UV radiation caused chlorophyll degradation and a decrease in the maximum quantum yield of photosystem II (PSII), together with an increase in the mechanisms of non-photochemical dissipation of energy (increase in the xanthophyll index). Cadmium was more stressing than UV radiation, since the metal also influenced photosynthesis globally and caused a decrease in net photosynthetic rates, in the effective quantum yield of photosynthetic energy conversion of PSII, and in the maximal apparent electron transport rate through PSII. Ultraviolet radiation increased the level of trans-p-coumaroylmalic acid and cadmium increased trans-phaselic and feruloylmalic acids. The increase in these compounds was probably related to both a more efficient absorption of harmful UV radiation and an enhanced protection against oxidative stress. DNA damage was specifically caused by UV-B radiation, but was intensified under the presence of cadmium, probably because the metal impairs the DNA enzymatic repair mechanisms. Ultraviolet radiation and cadmium seemed to operate additively on some physiological processes, while other responses were probably due to either factor alone.