Effects of drought stress on antioxidant enzyme, photosynthetic pigment and flavonoid pathway in two desert shrubs

Antioxidant enzyme activity, photosynthetic pigment content, and free malondialdehyde (MDA), as well as flavonoid content and the key enzyme activity in the flavonoid pathway were determined in two desert shrubs, Caryopteris mongolica Bunge and Reaumuria soongorica (Pall.) Maxim. under drought stress. The free MDA content was enhanced during the experimental period, which may be an indicator of oxidative stress. Superoxide dismutase (SOD), peroxidase (POD), and ascorbate peroxidase (APX) activities in C. mongholica showed a significant increase during the experiment, but catalase (CAT) activity was slightly decreased. On the other hand, POD and APX activities showed a significant increase and SOD and CAT activity data had no significant changes in R. soongorica. APX, SOD, and CAT activities were higher in R. soongorica than in C. mongholica, but MDA content was lower, indicating that the lower values of MDA were attributed to higher activities of antioxidant enzyme in R. soongorica. Chlorophyll content decreased significantly in the two shrubs during the experiment, which indicated that there was a photoprotection mechanism through reducing light absorbance by decreasing pigments content. Caretonoids content increased in C. mongholica and decreased in R. soongorica. The ratio of Chla/Chlb decreased significantly but caretonoids/Chl revealed a significant increase in the two shrubs, which could be explained as no decrease of peripheral light-harvesting complexes and a higher tolerance to drought. Total flavonoid content and the activities of phenylalanine ammonialyase (PAL) and chalcone isomerase (CHI) showed different changes between C. mongholica and R. soongorica after treatment. These values decreased in R. soongorica and increased in C. mongholica except for PAL activity. However, anthocyanin content increased in the two shrubs, indicating that there was a different regulation response in the flavonoid pathway in the two shrubs under drought stress, and anthocyanin should be an important antioxidant both in the shrubs. Our results demonstrated the different responses of antioxidant defense and drought tolerance ability between the two shrubs.     

Effects of salt and alkali stress on Reaumuria soongorica germination

Seed germination and early seedling growth are crucial stages for plant establishment. Two neutral (NaCl and Na₂SO₄) and two alkali (NaHCO₃ and Na₂CO₃) salts were selected to investigate their effects on germination and recovery responses in Reaumuria soongorica. Results show that both salt types significantly reduced germination and radicle elongation. The rate of germination and emergence of R. soongorica seeds continuously decreased as salinity increased, and the time to achieve maximum germination rate was delayed. The speed of seed germination dropped rapidly as salt concentration increased. Alkaline salts restricted the germination rate of R. soongorica seeds, and stresses resulting from alkaline salts and high concentrations of neutral salts resulted in many deformed seedlings. The length of the radicle and germ decreased with increasing salt concentration, but certain concentrations of salt and increased pH promoted germ growth. The results of regression analysis show that salt concentration was the dominant factor inhibiting R. soongorica seed germination rate. Salinity, buffering capacity, and pH all affected embryo growth, but salinity had the most pronounced effect. Seed viability under highly saline conditions appears to be a better indicator of adaptation to saline environments than seed germination under saline conditions.     

不同生境下红砂(Reaumuria soongorica)的生理生化特征及适应性

测定了荒漠地区超旱生小灌木红砂在3种生境(山前荒漠、山前戈壁和中游戈壁)下的叶片水分状态、抗氧化酶活性及渗透调节物质含量,以探讨荒漠植物在不同生境下的生理生化特征及对荒漠极端环境的适应机制。结果表明:在3种生境下,红砂叶片相对含水量和叶片水势随着土壤含水量的下降而下降,叶绿素含量随着土壤含水量的下降而升高。随着生境土壤干旱的加剧,超氧化物歧化酶(SOD)及过氧化物酶(POD)活性逐渐升高,而过氧化氢酶活性下降。在土壤含水量较少的戈壁生境,渗透调节物质含量上升。SOD和POD活性的升高是红砂抵御干旱环境的主要抗氧化保护机制;渗透调节在红砂叶片适应干旱胁迫的过程中发挥着重要作用,高的渗透调节能力使红砂在水分不足的条件下维持较低的渗透势,有利于植物吸水,从而增强其耐旱性。     

Adaptive evolution of rbcL in Reaumuria soongarica (Tamaricaceae)

In the field of phylogenetic analyses, the rbcL gene encoded large subunit Ribulose-1,5-biphosphate carboxylase/oxygenase (Rubisco, EC4.1.1.39), which plays a crucial role in the process of photosynthesis for most terrestrial plants, has been considered to be conserved; however, recent controversy regarding rbcL conservation has appeared since it was proposed to be under natural selection within all principal lineages of land plants. In this study, by examining the variation of DNA and protein sequences among 17 species in the family Tamaricaceae, three nonsynonymous mutations were identified to be under positive selection. The favored sites were located in the alph-helix domains of Rubisco, with decreased hydrophobicity and increased entropy, which could facilitate CO₂ penetration into the active site of Rubisco. We also found that the expression level of rbcL in different genotypes of Reaumuria soongarica shifted in response to various stresses such as drought, temperature, salt, and light. This study not only sheds light on the functional/structural features of Rubisco in the evolution scenarios from C₃-like into C₄ in Tamaricaceae but also provides useful information on directing genetic performance to enhance photosynthesis efficiency of desert plants for sustaining fragile desert ecosystems; furthermore, it promotes the ability to cope with desert aridification and global warming.     

Effects on antioxidant enzyme activities and osmolytes in Halocnemum strobilaceum under salt stress

The seedlings of Halocnermum strobilaceum were cultivated in 0.5% hoagland nutrient solution containing 0.0%, 0.9%, 2.7% and 5.4% of NaCl as well as composite salt (Na⁺, Ca²⁺, K⁺, Si⁴⁺) for 20 days; all the contents are in weight ratio. Succulent level, inorganic ions (Na⁺, K⁺), organics such as betaine, proline, malondialdehyde, and antioxidant enzyme activities including superoxide dismutase (SOD), catalase (CAT), peroxidase (POD), betaine aldehyde dehydrogenase (BADH) were measured to reveal its salt tolerance mechanism. When the composite salt concentration reaches 5.4%, SOD activity level, and MDA content is five times the control group; when it reaches 2.7%, the succulent level of seedlings, and the content of K⁺ in roots is nearly two times the NaCl treatment; the dry weight is more than three times the control group; with the NaCl treatment, MDA is three times the contrast; when the salt concentration is 2.7%, POD reaches the maximum. Results indicate that Si⁴⁺, K⁺, and Ca²⁺ from composite salt in the roots of H. strobilaceum improved the water-holding capacity. The activities of antioxidant enzyme were raised by the accumulation of proline and betaine, which increased the salt tolerance. The absorption of K⁺ promoted the high ratio of K⁺/Na⁺ and alleviated the damage of cell membranes of H. strobilaceum, which is associated with osmotic contents such as betaine and proline.     

Comparison of anatomical structure and photosynthetic characteristics between the two photosynthetic organs of the desert plant Hedysarum scoparium

The desert plant Hedysarum scoparium uses leaflets and rachises as its photosynthetic organs. The abundance of leaflets was lower under unfavorable environmental conditions and higher with improved water conditions. To examine the characteristics associated with the adaptation of H. scoparium to its environment, we selected plants with both compound leaves and rachis without leaflets to study the anatomical structures and gas exchange characteristics of the two organs. The results show that the water storage tissues in rachises were more developed compared with the leaflets. The diurnal courses of the net photosynthetic rate for the rachis and the leaflet were both in a bimodal pattern. Meanwhile, both two peak values of the rachis were significantly higher than those of the leaflet. The daily average transpiration rate was significantly higher in the rachis than in the leaflet in order to lower the temperature of the rachises. It was concluded that under desert drought conditions, the leaflets of H. scoparium were partially or completely degraded to reduce the transpiration area as an adaptive response to water deficit, and only the rachises were retained as photosynthetic organ. The rachises were found to be better suited to a desert habitat than the leaflets.     

Responses to salt stress in the halophyte Plantago crassifolia (Plantaginaceae)

General responses to salt stress have been investigated in the halophyte Plantago crassifolia. Seed germination was strongly inhibited by NaCl, although seed viability and germination capacity were not affected by salt pre-treatments. A concentration-dependent inhibition of plant growth was observed in the presence of NaCl, which was accompanied by the accumulation of Na⁺ ions in the leaves, as determined by cation exchange HPLC. A 20-fold increase of proline content in leaves was observed when plants were treated with 500 m NaCl, suggesting a protective role against high salinity stress for this amino acid, whose possible mechanism of action is discussed.     

Physiological responses and adjustment mechanisms of the dominate species of natural vegetation of Eastern Tengger Desert

Reaumuria soongorica and Salsola passerina have significant differences in ecophysiological characteristics, which change with the environmental variations. Although they live together for a long period of time, their adaptive mechanisms to environmental stresses are very different. As two extreme xerophytes, Reaumuria soongorica and Salsola passerina differ significantly from other psammophytes in ecophysiological characteristics; they can survive in lower water potential, and can even grow in piedmont areas. Low water potential may be related to the existence of osmosis-regulating substance, such as praline, which can strengthen the capacity of water absorption. Compared to other psammophytes, Reaumuria soongorica and Salsola passerina have a higher degree of photo-inhibition under the same condition, and the photo-inhibition can lead to destruction of the photosynthetic pigment, nevertheless, this photo-inhibition can be repaired under suitable conditions in the morning and evening.     

Eco-physiological mechanisms of Caragana korshinskii Kom. adaptation to extreme drought stress: Leaf abscission and maintaining stem chloroplast integrity

In order to study the eco-physiological mechanisms of C. korshinskii adaptation to extreme drought stress, we investigated the variations of water content in soil, leaves, and stems, the chlorophyll a and b and the carotenoid content in leaves and stems, as well as changes of chloroplast ultrastructure in 2-year-old C. korshinskii specimens during a progressive soil drought process (by ceasing watering until all leaves were shed) and a subsequent rehydration process. During the dehydration process, the chlorophyll a and b and carotenoid contents in the leaves decreased, as did the carotenoid content in the stems. During the 4-day rehydration process, the chlorophyll a and b and carotenoid contents in the leaves and stems increased and gradually returned to normal levels. During ongoing drought stress, chloroplasts in the leaves broke away from cell walls and appeared in the center of cells. Under severe drought stress, the mesophyll ultrastructure and chloroplast configuration in leaves were irreversibly disturbed, as manifested by the inner and outer membranes being destroyed; the thylakoid system disintegrated, the starch grain disappeared, and parts of cell tissue were dismantled into debris. However, the mesophyll ultrastructure and chloroplast configuration in the stems remained complete. This indicates that C. korshinskii utilizes leaf abscission to reduce the surface area to avoid damage from extreme drought stress, and maintains chloroplast integrity and a considerable amount of chlorophyll to enable a rapid recovery of photosynthesis under the rehydration process.     

Display-sites selection by houbara bustard (Chlamydotis [undulata] macqueenii) in Mori, Xinjiang, People''s Republic of China

Display-site selection by houbara bustards was studied in Mori, Xinjiang, China during the breeding seasons from April to June 2000. Most of the habitats chosen for displaying were short sub-shrubby and open areas close to high shrub patches. The displaying males clearly prefer low covered areas and avoid densely covered and high vegetation sites. The vegetation density and number of plant species at display sites were significantly lower from that at randomly selected sites. The average distance to the closest shrub patch was significantly shorter at display sites than at random sites. Plant species richness, vegetative density, vegetative cover and distance to the shrub patches are possibly the most important factors that determined the display-sites selection of houbara bustard.     

Germination of Callitris seeds in relation to temperature, water stress, priming, and hydration–dehydration cycles

The effects of temperature, water stress, hydration–dehydration cycles and seed priming on the germination of Callitris verrucosa and Callitris preissii, two Australian semi-arid coniferous tree species, were investigated. Optimum germination occurred at 18°C, with a minimum germination time of 8–9 days for both species. At this temperature, germination was inhibited at osmotic potentials lower than −1·0 MPa, but the capacity to germinate at low osmotic stress increased as the temperature decreased. Seed priming and hydration–dehydration cycles did not reduce seed viability, and Callitris seeds appear to retain the physiological changes induced by short-term hydration, as the time to the onset of germination was decreased to about 3 days. The capacity of Callitris seeds for incremental germination is likely to increase overall germination success in a low rainfall environment.     

Growth response of the saltbush Atriplex nummularia L. to inoculation with the arbuscular mycorrhizal fungus Glomus intraradices

Plantlets of Atriplex nummularia were inoculated with the arbuscular mycorrhizal fungi Glomus intraradices in a pot experiment. Plants were grown in a low P soil. Highly significant growth response of a Chenopodiaceae was recorded for the first time. Mycorrhizal colonization of roots was well developed, internal hyphae and vesicles were observed, but not arbuscules. These observations suggest that arbuscules are not necessary to obtain significant growth stimulation from mycorrhizal inoculation. Atriplex nummularia is already used as forage crops, its high mycorrhizal dependency offers possibilities to develop this production and revegetation strategies.     

The diet of the Long-eared Owls, Asio otus, in the desert of northwest China

The contents of regurgitated Long-eared Owls (Asio otus) pellets collected in Minqin Desert Experimental Research Station of northwestern China were analysed. A total of 303 individuals representing five species of rodents and one of bird were identified in the pellets. By frequency, the most common species taken by Long-eared Owls was Meriones meridianus, followed by Cricetulus barabensis and Phodopus roborovskii. By biomass, Meriones meridianus was the most important species taken, followed by C. barabensis and Rhombomys opimus. Our results suggest that the Long-eared Owls were nocturnal hunters and fed on a wide range of animals with respect to their habitats.     

Photosynthetic responses of Larrea tridentata to a step-increase in atmospheric CO2at the Nevada Desert FACE Facility

Of all terrestrial ecosystems, the productivity of deserts has been suggested to be the most responsive to increasing atmospheric CO₂. The extent to which this prediction holds will depend in part on plant responses to elevated CO₂under the highly variable conditions characteristic of arid regions. The photosynthetic responses ofLarrea tridentata , an evergreen shrub, to a step-increase in atmospheric CO₂(to 550 μmolmol⁻¹) were examined in the field using Free-Air CO₂Enrichment (FACE) under seasonally varying moisture conditions. Elevated CO₂substantially increased net assimilation rate (A_net) in Larrea during both moist and dry periods of the potential growing season, while stomatal conductance (g_s) did not differ between elevated and ambient CO₂treatments. Seasonal and diurnal gas exchange dynamics in elevated CO₂mirrored patterns in ambient CO₂, indicating that elevated CO₂did not extend photosynthetic activity longer into the dry season or during more stressful times of the day. Net assimilation vs. internal CO₂(A/C_i) responses showed no evidence of photosynthetic down-regulation during the dry season. In contrast, after significant autumn rains, A_max(the CO₂saturated rate of photosynthesis) and CE (carboxylation efficiency) were lower in Larrea under elevated CO₂. In situ chlorophyll fluorescence estimation ofLarrea Photosystem II efficiency (F_v/F_m) responded more to water limitation than to elevated CO₂. These findings suggest that predictions regarding desert plant responses to elevated CO₂should account for seasonal patterns of photosynthetic regulatory responses, which may vary across species and plant functional types.     

Carbon accumulation, nitrogen and phosphorus use efficiency of Sophora davidii seedlings in response to nitrogen supply and water stress

A greenhouse experiment was conducted to explore whether additional nitrogen (N) supply could enhance carbon (C) accumulation, and phosphorus (P) use efficiency (NUE_P) of Sophora davidii seedlings under dry conditions. Two-month-old seedlings were subjected to a completely randomized design with three water (80, 40 and 20% water field capacity (FC)) and three N supply (N0: 0, Nl: 92 and Nh: 184 mg N kg⁻¹ soil) regimes. Water stress decreased C, N and P accumulation, NUE_P, N and P uptake efficiency (NUtE_N and NUtE_P) regardless of N supply. The S. davidii seedlings exhibited strong responses to N supply, but the responses were not consistent with the various N supply levels. Nl increased C, N and P accumulation, and improved NUE_P, NUtE_N and NUtE_P in the same water treatment. In contrast, Nh did few or even depress effects on C, N and P accumulation, and NUE_P, although NUtE_N and NUtE_P increased with Nh in the same water treatment. Even so, NUE_N decreased with increase of N supply in the same water treatment. The results suggested that appropriate or low N supply should be recommended for S. davidii seedling establishment in dry environment by improving C accumulation and NUE_P.     

Foliar carbohydrate differs between Picea crassifolia and Sabina przewalskii with the altitudinal variation of Qilian Mountains, China

Nonstructural carbohydrates(NSC) and nitrogen metabolism strongly influence growth and development in plants. The biosynthesis of cellulose and lignin(structural carbohydrates, SC) depends largely on the supply of NSC. We desire to examine which hypothesis, carbon limitation or growth limitation, best fits the alpine plant response between NSC, SC, carbon(C), nitrogen(N) and altitude. We compared the foliar concentrations of carbohydrates, C and N between the leaves of Picea crassifolia(lower-elevation tree-line species) and Sabina przewalskii(high-elevation tree-line species) in their response to changing elevation. Our site was located in the mid-northern area of Qilian Mountains, China. We found that foliar soluble sugar(SG) concentrations were significantly higher in P. crassifolia than in S. przewalskii at the 2,700–3,400 m level. Foliar NSC levels in P. crassifolia increased at the 2,700–3,100 m level, indicating that growth was limited gradually resulting in a surplus of NSC(to conform to GLH), subsequently decreasing at the 3,100–3,400 m level, the assimilation declined leading to C deficit(to conform to CLH). SC(SC metabolism disorders at 3,100–3,400 m), C, N and starch were significantly lower in P. crassifolia than in S. przewalskii. Conversely, foliar SG concentration shows a fall-rise trend with increasing elevation for S. przewalskii. SC concentration in S. przewalskii leaves decreased with an increase of elevation and has a significantly positive correlation to N concentration marking the assimilation of plants. Therefore, the high-elevation tree-line species(S. przewalskii) utilize or store more foliar SG leading to a decrease of SG concentration for survival and growth/regrowth in an adverse environment, higher total C, N, SC, starch contents and lower NSC level. Also, their change trends along the elevational gradient in leaves of S. przewalskii indicate better assimilation strategies for SG use under environmental stress compared to P. crassifolia. This indicates that foliar C metabolism along the elevation follows the principle of the growth-limitation hypothesis(GLH) or carbon limitation hypothesis(CLH), which depends on the acclimation of different alpine life-forms to the environment.     

Climatic response of Pinus sylvestris var. mongolica tree-ring width and precipitation reconstruction for the northern Greater Higgnan Mountains, China, since 1720

August-June precipitation has been reconstructed back to AD 1720 for the northern Greater Higgnan Mountains, China, by use ofPinus sylvestris var. mongolica tree-ring width. The reconstruction explains the variance of 39% in observed precipitation from 1973 to 2008. Some extremely dry/wet signals in historical documents and other precipitation reconstructions in previous studies are precisely captured in our reconstruction. Wet periods occurred during the periods of 1730 to 1746, 1789 to 1812, 1844 to 1903, 1920 to 1930, 1942 to 1961, and 1985 to 1998; while periods of 1747 to 1788, 1813 to 1843, 1904 to 1919, 1931 to 1941, and 1962 to 1984 were relatively dry. Power spectral and wavelet analyses demon- strated the existence of significant 24-year, 12-year, and 2-year cycles of variability.