Low induction of non‐photochemical quenching and high photochemical efficiency in the annual desert plant Anastatica hierochuntica

Non‐photochemical quenching (NPQ) plays a major role in photoprotection. Anastatica hierochuntica is an annual desert plant found in hot deserts. We compared A. hierochuntica to three other different species: Arabidopsis thaliana, Eutrema salsugineum and Helianthus annuus, which have different NPQ and photosynthetic capacities. Anastatica hierochuntica plants had very different induction kinetics of NPQ and, to a lesser extent, of photosystem II electron transport rate (PSII ETR), in comparison to all other plants species in the experiments. The major components of the unusual photosynthetic and photoprotective response in A. hierochuntica were: (1) Low NPQ at the beginning of the light period, at various light intensities and CO₂ concentrations. The described low NPQ cannot be explained by low leaf absorbance or by low energy distribution to PSII, but was related to the de‐epoxidation state of xanthophylls. (2) Relatively high PSII ETR at various CO₂ concentrations in correlation with low NPQ. PSII ETR responded positively to the increase of CO₂ concentrations. At low CO₂ concentrations PSII ETR was mostly O₂ dependent. At moderate and high CO₂ concentrations the high PSII ETR in A. hierochuntica was accompanied by relatively high CO₂ assimilation rates. We suggest that A. hierochuntica have an uncommon NPQ and PSII ETR response. These responses in A. hierochuntica might represent an adaptation to the short growing season of an annual desert plant.     

Melanogenesis inhibitors from the desert plant Anastatica hierochuntica in B16 melanoma cells

The methanolic extract from the whole plants of Anastatica hierochuntica, an Egyptian herbal medicine, was found to inhibit melanogenesis in theophylline-stimulated murine B16 melanoma 4A5 cells. Among the constituents isolated, anastatin A, silybin A, isosilybins A and B, eriodictyol, luteolin, kaempferol, quercetin, hierochins A and B, (2R,3S)-2,3-dihydro-2-(3,4-dimethoxyphenyl)-3-hydroxymethyl-5-(2-formylvinyl)-7-hydroxybenzofuran, (+)-dehydrodiconiferyl alcohol, (+)-balanophonin, 1-(4-hydroxy-3-methoxyphenyl)-2-{4-[(E)-3-hydroxy-1-propenyl]-2-methoxyphenoxy}-1,3-propanediol, and 3,4-dihydroxybenzaldehyde substantially inhibited melanogenesis with IC₅₀ values of 6.1–32 μM. With regard to the mechanism of action of silybins and isosilybins, the inhibition of tyrosinase activity suggested to be important. In addition, isosilybins A and B inhibited the mRNA expression of TRP-2, but silybins A and B oppositely enhanced the mRNA expression of tyrosinase and TRP-1 and -2 at 10 and/or 30 μM, and the inhibition of phosphorylation of extracellular signal-regulated kinases (ERK1/2) is involved in the enhanced expression of mRNA, at least in part, similar to that of PD98059.     

Anastatins A and B,new skeletal flavonoids with hepatoprotective activities from the desert plant Anastatica hierochuntica

New skeletal flavonoids, anastatins A and B, were isolated from the methanolic extract of an Egyptian medicinal herb, the whole plants of Anastatica hierochuntica. Their flavanone structures having a benzofuran moiety were determined on the basis of chemical and physicochemical evidence. Anastatins A and B were found to show hepatoprotective effects on D-galactosamine-induced cytotoxicity in primary cultured mouse hepatocytes and their activities were stronger than those of related flavonoids and commercial silybin.     

Anatomical significance of the hygrochastic movement in Anastatica hierochuntica

BACKGROUND AND AIMS: Unlike the dispersal mechanisms of many desert plants, the whole dead skeleton of Anastatica hierochuntica is involved in seed dispersal and preservation. This process depends on the hygrochastic nature of the lignified conducting tissue that bends when dry and straightens under wet conditions. An anatomical interpretation of this mechanical movement was investigated. METHODS: An anatomical study of the stem was conducted on the juvenile plants raised under different water treatments and on the branch-orders of adult A. hierochuntica size-classes. KEY RESULTS: In the juvenile stem of A. hierochuntica, the area of cortex, conducting tissue and pith increased with water availability. However, the hydraulic conductance decreased, resulting in a better withdrawal of water in water-stressed plants. The anatomical investigation of the hygrochastic mechanism revealed an asymmetric distribution of the cortical tissues, with the conducting tissues of the stem of juvenile and adult plants being larger in the lower side. The hydraulic conductance was better in the basal and middle branch-orders than the terminal ones, permitting better conductance of water to the subsequent branch-orders. CONCLUSIONS: The lignified conducting tissue of the whole stem, having a hygrochastic nature, controls the movement of the branches. The greater amount of conducting tissue associated with a higher density of wide xylem vessels was observed in the lower side of the stem as compared with the upper side. Consequently, the conducting tissue in the lower side of the stem was suggested to be more effective in the opening process of the curled dry branches through better and more rapid conductance of water. Alternatively, due to the few narrow xylem vessels in the upper side of the stem, it was likely that the conducting tissue in the upper side is more effective in the closing process by providing more rapid drying. The mechanical rise of water and the related hygrochastic efficiency were maximized in the basal and middle branch-orders that are mostly involved in the mechanical movement.     

Photosynthesis and photoprotection in mangroves under field conditions

Net CO₂ exchange and in vivo chlorophyll fluorescence were studied in mangrove (Rhizophora stylosa) leaves at a field site in Western Australia, and leaf samples were collected for the analysis of enzymes and substrates potentially involved in anti-oxidant photoprotection. Photosynthesis saturated at 900 μmol quanta m^?2 s^?1 and at no more than 7.5 μmol CO₂ m^?2 s^?1. However, fluorescence analysis indicated no chronic photoinhibition: F_v:F_m was 0.8 shortly after sunset, and quantum efficiencies of PSII were high up to 500 μmol quanta m^?2 s^?1. Electron flow through PSII was more than 3 times higher than electron consumption through Calvin cycle activity, however, even with photorespiration and temperature-dependent Rubisco specificities taken into account. Acknowledging the growing body of literature attributing a role to antioxidant systems in photoprotection, we also assayed the activities of superoxide dismutase (SOD) and several enzymes potentially involved in H₂O₂ metabolism. Their levels of maximal potential activity were compared with those in greenhouse-grown mangroves (R. mangle), and growth chamber-grown peas. Monodehydroascorbate reductase activities were similar in all species, and glutathione reductase was lower, and ascorbate peroxidase ～40% higher, in the mangroves. SOD activities in field-grown mangroves were more than 40 times those in peas. Our results support the hypothesis that O₂ may be a significant sink for photochemically derived electrons under field conditions, and suggest an important role for O₂^? scavenging in photoprotection. However, when relative patterns are compared between species, imbalances between SOD and the other enzymes in the mangroves suggest that more components of the system (e.g. phenolics or peroxidases) are yet to be identified.     

Drought tolerance of germinating seeds and young seedlings of Anastatica hierochuntica L.

Summary Drought tolerance of germinating seeds and young seedlings of Anastatica hierochuntica L. was studied under controlled conditions. When seeds at various stages of germination were air-dried, or alternatively were desiccated over CaCl₂ to the same or a lower water level than in the air-dried seeds, and after various periods of storage were rewetted, they readily continued germination or growth. Similarly, 60% of dried young seedlings with rootlets 4–6 mm long that had been dehydrated for one week, renewed their growth 8 h after wetting, their shrivelled, dried rootlets imbibing water. Recovery was adversely affected by both longer periods of germination prior to the dehydration, or by the longer duration of storage of the redried seeds/seedlings. The higher rate of dehydration also reduced recovery. It is argued that despite this tolerance of seedlings to drought, the Rose of Jericho cannot be considered a resurrection plant. Compared with other desert annuals, this behavior is specific for Anastatica. However, it may occur also in other annual species of populations sequestered within the driest of desert habitats.     

Solar tracking response to drought in a desert annual

Summary The responses to drought of the solar tracking winter annualLupinus arizonicus (Wats.) were examined under field and laboratory growth regimes. Under drought conditions tracking movements were maintained until the plant reached the wilting point. The leaves and leaflets were observed to cup in response to decreases in the xylem water potential. This resulted in a negative, but linear relationship between the cosine of the angle of incidence of the direct solar beam upon the leaf and water potential. Leaf conductance was also significantly related to xylem water potential in natural populations. Leaf and leaflet cupping seem to be responding directly to changing plant water status, resulting in reduced solar radiation load to drought stressed plants. It is hypothesized that this property may have adaptive consequences in relation to the water relations, energy budget, and carbon balance of the plant.     

Photosynthetic regulation of C4 desert plant Haloxylon ammodendron under drought stress

About 20-year-old desert plants of C₄ species, Haloxylon ammodendron, growing at the southern edge of the Badain Jaran Desert in China, were selected to study the photosynthetic characteristics and changes in chlorophyll fluorescence when plants were subject to a normal arid environment (AE), moist atmospheric conditions during post-rain (PR), and the artificial supplement of soil water (SW). Results showed that under high radiation, in the AE, the species down-regulated its net assimilation rate (A) and maximum photochemical efficiency of PS II (Fv/Fm), indicating photoinhibition. However, under the PR and SW environments, A was up-regulated, with a unimodal diurnal course of A and a small diurnal change in Fv/Fm, suggesting no photoinhibition. When the air humidity or SW content was increased, the light compensation points were reduced; light saturation points were enhanced; while light saturated rate of CO₂ assimilation (A _max) and apparent quantum yield of CO₂ assimilation (Φ_C) increased. Φ_C was higher while the A _max was reduced under PR relative to the SW treatment. It was concluded that under high-radiation conditions drought stress causes photoinhibition of H. ammodendron. Increasing air humidity or soil moisture content can reduce photoinhibition and increase the efficiency of solar energy use.     

Photosynthesis and photoprotection responses to water stress in the wild-extinct plant Lysimachia minoricensis

Lysimachia minoricensis is an endemic species of the Balearic Islands that has become extinct in the wild, but persists in botanical gardens. Attempts of re-introducing the species into its natural habitat, which consisted in temporary dry streams, have failed. Low genetic variability has been reported for the garden individuals, suggesting that a reduced potential to adapt to environmental changes could be among the reasons for its extinction. In the present study, we particularly test whether photosynthesis and photoprotection responses of this species to water stress could help explaining the lack of success of this species in its natural habitat.

Plants of L. minoricensis were grown in pots in a growth chamber. Soil water depletion was imposed over 20 days by stopping irrigation. Early stomata closure was observed in response to soil water depletion while leaves desiccated progressively. Although net photosynthesis was low in irrigated plants, due to a remarkably low mesophyll conductance to CO₂, substantial photosynthetic activity was kept at severe drought, where leaf relative water content was as low as 50%, suggesting that L. minoricensis is a very drought-tolerant species. In parallel with decreased photosynthesis, thermal dissipation of the excess light and photorespiration progressively increased. The former was linearly related to increased de-epoxidation of the xanthophylls cycle. Photoprotection was effective, as pre-dawn maximum photochemical efficiency was maintained higher than 0.75 through the entire experiment. Moreover, photosynthetic capacity was largely (80%) recovered only 24 h after re-watering. These results show that stomatal regulation, photosynthetic metabolism and photoprotection in L. minoricensis are well adapted to water stress, suggesting that additional factors may be responsible for its status as a wild-extinct plant.     

Water response of the hygrochastic skeletons of the true rose of Jericho (Anastatica hierochuntica L.)

Summary The hygrochastic apparatus in Anastatica hierochuntica was investigated from the standpoint of morphology of the skeleton and fruit and its contribution to the regulation of seed dispersal in accordance with the water conditions.Water absorption by the dry skeletons upon shoot immersion under laboratory conditions is a rapid process requiring only a few minutes. It is followed by a slow opening of the curled branches of the skeleton. Maximal opening is attained by the saturated skeletons in approximately 2h. Absorption of water and subsequent opening occur also when only the main root of the skeleton is immersed in water. The process was duplicated also by intact skeletons irrigated in situ. Skeletons closure in response to evaporation is initially slow, noticable curling of the branches obtained only after 60% of the water is lost from the saturated skeletons. Opening of the skeleton exposes most of the fruits to the force of falling raindrops. The fact that no spontaneous opening of fruits occurred under most laboratory procedures of skeleton immersion suggests that in nature two successive processes occur: the skeletons must first uncurl-hygrochasticly so that raindrops can impinge forcefully on the appendages of the fruit valves and thus cause seed release by ombrohydrochory. Under adequate rainfall, the first fruits to open are those at the outer infructescences; heavier rainfall will release seeds from the upper fruits of the inner infructescences, and later, from fruits at their bases. The force that binds the valves of the fruits to the septa is variable, its strength correlated with different structure of the fruist at different positions on the skeleton. This ensures a rate of seed release commensurate with the rainfall, and secures at least a minimum of water for the first seeds. The heterogeneity of skeleton populations insofar as age of the plants is concerned, further ensures that seed dispersal in a given area will be proportional to the amount of rainfall.     

Photosynthesis, photoprotection, and growth of shade-tolerant tropical tree seedlings under full sunlight

High solar radiation in the tropics is known to cause transient reduction in photosystem II (PSII) efficiency and CO(2) assimilation in sun-exposed leaves, but little is known how these responses affect the actual growth performance of tropical plants. The present study addresses this question. Seedlings of five woody neotropical forest species were cultivated under full sunlight and shaded conditions. In full sunlight, strong photoinhibition of PSII at midday was documented for the late-successional tree species Ormosia macrocalyx and Tetragastris panamensis and the understory/forest gap species, Piper reticulatum. In leaves of O. macrocalyx, PSII inhibition was accompanied by substantial midday depression of net CO(2) assimilation. Leaves of all species had increased pools of violaxanthin-cycle pigments. Other features of photoacclimation, such as increased Chl a/b ratio and contents of lutein, β-carotene and tocopherol varied. High light caused strong increase of tocopherol in leaves of T. panamensis and another late-successional species, Virola surinamensis. O. macrocalyx had low contents of tocopherol and UV-absorbing substances. Under full sunlight, biomass accumulation was not reduced in seedlings of T. panamensis, P. reticulatum, and V. surinamensis, but O. macrocalyx exhibited substantial growth inhibition. In the highly shade-tolerant understory species Psychotria marginata, full sunlight caused strongly reduced growth of most individuals. However, some plants showed relatively high growth rates under full sun approaching those of seedlings at 40?% ambient irradiance. It is concluded that shade-tolerant tropical tree seedlings can achieve efficient photoacclimation and high growth rates in full sunlight.     

荒漠一年生植物小画眉草的种群动态调节与模拟

以分布在沙波头沙漠试验研究站人工固沙植被中的一年生植物小画眉草为例,研究了荒漠一年生植物不同时间尺度上的种群动态与调节机制,试验结果表明：（1）一年生植物小画眉草种群的续存与其种子库中种子的分批萌发对策密切相关;（2）同一个生长季内,制约小画眉草种群数量变化的关键因子随降水条件的不同而变化,即当环境适宜度较小时（干旱胁迫）,非生物因子（降水）限制种群数量;当环境适宜度较大时,密度依赖的竞争作用调节种群大小;（3）在较长的生态时间尺度上土壤特性（结皮厚度,养分含量）也是影响小画眉草种群动态的一个重要因素;（4）小画眉草种群的存活曲线属于C型,说明该种是较为典型的r－对策者。     

Evolutionary and plastic changes in a native annual plant after a historic drought

Severe droughts are forecast to increase with global change. Approaches that enable the study of contemporary evolution, such as resurrection studies, are valuable for providing insights into the responses of populations to global change. In this study, we used a resurrection approach to study the evolution of the California native Leptosiphon bicolor (true babystars, Polemoniaceae) across populations differing in precipitation in response to the state's recent prolonged drought (2011–2017). In the Mediterranean climate region in which L. bicolor grows, this historic drought effectively shortened its growing season. We used seeds collected both before and after this drought from three populations found along a moisture availability gradient to assess contemporary evolution in a common garden greenhouse study. We coupled this with a drought experiment to examine plasticity. We found evolution toward earlier flowering after the historic drought in the wettest of the three populations, while plasticity to experimental drought was observed across all three. We also observed trade‐offs associated with earlier flowering. In the driest population, plants that flowered earlier had lower intrinsic water‐use efficiency than those flowering later, which was an expected pattern. Unexpectedly, earlier flowering plants had larger flowers. Two populations exhibited evolution and plasticity toward smaller flowers with drought. The third exhibited evolution toward larger flowers, but displayed no plasticity. Our results provide valuable insights into differences among native plant populations in response to drought.     

Photosynthesis and photoprotection in Nicotiana tabacum L. in vitro-grown plantlets

Nicotiana tabacum L. plantlets were cultured in vitro photoautotrophically (0% sucrose) and photomixotrophically (3% or 5% sucrose) at two irradiances (80 or 380 mumol m-2 s-1) with the aim of investigating the effect of these culture conditions on photosynthetic parameters and on protective systems against excess excitation energy. In plantlets grown photoautotrophically under higher irradiance photoinhibition was demonstrated. These plantlets had a decreased chlorophyll (Chl) a + b content and Chl a/b ratio, an increased content of xanthrophyll cycle pigments and a higher deepoxidation state, a decreased maximum photochemical efficiency of photosystem II (PS II) and actual photochemical efficiency of PS II, and an increased non-photochemical quenching. In the photoautotrophically grown plantlets and those photomixotrophically grown with 3% sucrose, the increase of growth irradiance from 80 to 380 mumol m-2 s-1 stimulated the activities of ascorbate-glutathione cycle enzymes with the exception of ascorbate peroxidase. Ascorbate peroxidase activity was not affected by the increase in growth irradiance but a significant decrease with increasing sucrose concentration was evident. The higher concentration of sucrose in the medium (5%) in combination with the higher irradiance inhibited photosynthesis (decrease in Chl a + b content and net photosynthetic rate) but no significant changes in activities of ascorbate-glutathione cycle enzymes were found. These results suggest that exogenous sucrose added to the medium improved high irradiance and oxidative stress resistance of the plantlets but the effect of sucrose is concentration dependent.     

The dynamics of abundance and incidence of annual plant species during colonization in a desert

We studied colonization of annual plants in small-scale disturbances and undisturbed soil for four years in the northern Negev desert. The experiment consisted of 24 patches of I m²: eight undisturbed patches, eight 30 cm deep pits and eight 20 cm high mounds. Disturbance removed the seed bank from the pits and mounds. Rainfall was average during 1992 and 1993 (163.5 and 157.0 mm), very low in 1994 (97.5 mm) and very high in 1995 (283.0 mm). For all ca 100 species in the assemblage, we measured abundance, as the average local density in patches occupied by the species, and incidence, the proportion of patches occupied. Abundance and incidence were positively correlated in the four years of the study (R^2-= 0.71, 0.56. 0. 56 and 0.64). The relationship became steeper during the course of colonization. A minority of species colonized all patches rapidly and increased exponentially in abundance. These high incidence and abundance species responded little or not at all to variations in rainfall. Most of the species experienced frequent local extinctions and colonizations in few patches. These low abundance and low to medium incidence species responded strongly to rainfall by fluctuations in abundance. Differences per species between abundance and/or incidence in pits, mounds and matrix were evident in many species along the entire relationship. These were mostly in favor of pits. We propose that the positions and trajectories of the individual species in the abundance-incidence phase plane signify constraints on population growth along the abundance axis, and on colonization and population persistence along the incidence axis. Since species with lower incidence and abundance respond strongly to rainfall variation, we conclude that their local populations and spatial distribution are to a large extent site-limited. High-abundance and high-incidence species have few or weak constraints and experience density-independent population growth, which is only seed-limited. The causes of site limitation and its effect on population growth and on eotonization and extinction processes differ for different species. In a number of species, abundance and incidence may be limited by dispersal, in conjunction with dispersal mode and patch-specific seed capture. Based on our case study we suggest that the trajectories of the species in the abundance-incidence phase plane are a useful tool for investigating assemblage dynamics.     

干旱下植物气孔运动的调控

概述了植物气孔对大气干旱和土壤干旱的反应,认为植物气孔对大气干旱的反应并不是一种反馈机制;并就干旱条件下植物气孔运动的水力学和化学信号调控机制进行了简要论述,认为虽然化学信号调控干旱下气孔运动更为广泛,但ABA不是唯一的化学信号,水分关系影响了信号的产生、运转和气孔对信号的敏感性,干旱条件下水力学和化学信号共同调控着植物的气孔运动。     

Photosynthesis, respiration and antioxidant enzymes in pepper leaves under drought and heat stresses

Leaf gas exchange, chlorophyll fluorescence and activities of antioxidant enzymes were studied in two pepper (Capsicum annuum L.) cultivars grown under drought (D) and heat (H), as well as under drought and heat in combination (HD). The drought-tolerant cv. Zhengjiao 13 exhibited greater net photosynthetic rate (P_N) and cytochrome respiratory pathway activity (R_SHAM), and lower contents of superoxide radical and hydrogen peroxide, as compared to the drought-sensitive cv. Longkouzaojiao. The P_N and R_SHAM decreased and ROS production increased under D and HD in both cultivars. As compared to the Longkouzaojiao, Zhengjiao 13 retained higher non-photochemical quenching (NPQ), photorespiration rate (R_L), and alternative respiratory pathways (R_KCN) under D and HD. Drought increased the superoxide dismutase (SOD) and ascorbate peroxidase (APX) activities in the cytosol, chloroplasts and mitochondria in the two cultivars. Although SOD and APX activities decreased in Longkouzaojiao under HD, SOD activity increased in Zhengjiao 13. There was no H-induced reactive oxygen species production due to increase of R_L, NPQ, R_SHAM, R_KCN and activities of SOD and APX. However, H slightly decreased the P_N. The results indicated that HD was more detrimental than either stress alone.