Growth and physiological responses of Agriophyllum squarrosum to sand burial stress

Agriophyllum squarrosum is an annual desert plant widely distributed on mobile and semi-mobile dunes in all the sandy deserts of China. We studied the growth and physiological properties of A. squarrosum seedlings under different sand burial depths in 2010 and 2011 at Horqin Sandy Land, Inner Mongolia to understand the ability and mechanism that A. squarrosum withstands sand burial. The results showed that A. squarrosum had a strong ability to withstand sand burial. Its survival rate, plant height and biomass increased significantly at a burial depth 25% of seedling height and decreased significantly only when the burial depth exceeded the height of the seedlings; some plants still survived even if the burial depth reached 266% of a seedling height. The malondialdehyde （MDA） content and membrane permeability of the plant did not change significantly as long as the burial depth was not greater than the seedling height; lipid peroxidation increased and cell membranes were damaged if the burial depth was increased further. When subjected to sand burial stress, superoxide dismutase （SOD） and peroxidase （POD） activities and free proline content increased in the seedlings, while the catalase （CAT） activity and soluble sugar content decreased. Sand burial did not lead to water stress. Reductions in photosynthetic area and cell membrane damage caused by sand burial may be the major mechanisms increasing mortality and inhibiting growth of the seedling. But the increases in SOD and POD activities and proline content must play a certain role in reducing sand burial damage.

Accumulation of soil organic carbon during natural restoration of desertified grassland in China's Horqin Sandy Land

China＇s Horqin Sandy Land,a formerly lush grassland,has experienced extensive desertification that caused considerable carbon（C） losses from the plant-soil system.Natural restoration through grazing exclusion is a widely suggested option to sequester C and to restore degraded land.In a desertified grassland,we investigated the C accumulation in the total and light fractions of the soil organic matter from 2005 to 2013 during natural restoration.To a depth of 20 cm,the light fraction organic carbon（LFOC） storage increased by 221 g C/m2（84%） and the total soil organic carbon（SOC） storage increased by 435 g C/m2（55%）.The light fraction dry matter content represented a small proportion of the total soil mass（ranging from 0.74% in 2005 to 1.39% in 2013）,but the proportion of total SOC storage accounted for by LFOC was remarkable（ranging from 33% to 40%）.The C sequestration averaged 28 g C/（m2·a） for LFOC and 54 g C/（m2·a） for total SOC.The total SOC was strongly and significantly positively linearly related to the light fraction dry matter content and the proportions of fine sand and silt＋clay.The light fraction organic matter played a major role in total SOC sequestration.Our results suggest that grazing exclusion can restore desertified grassland and has a high potential for sequestering SOC in the semiarid Horqin Sandy Land.

Characteristics of plant calcium fractions for 25 species in Tengger Desert

Little attention has been paid to plant calcium fractions in the desert.To address the characteristic of the calcium fractions of desert plants,we collected 25 plant species in Tengger Desert,observed the calcium crystals using an optical microscope and determined water soluble calcium,acetic acid soluble calcium,and hydrochloric acid soluble calcium.To do so,we used sequential fractionation procedures to probe the relationships among different functional groups,different growth forms,or different successional stages.The results showed that the psammophyte,the late successional plants,and the drought-resistant shrub and semi-shrub all held considerable calcium oxalate crystal compared to the grassland plants,the early successional plants,and the perennial herb.With the proceeding succession,the acetic acid soluble calcium decreased gradually,and the hydrochloric acid soluble calcium increased gradually.The perennial herb had more water soluble calcium,while shrub held greater hydrochloric acid soluble calcium.The grassland plants held more water soluble calcium,while psammophyte had greater hydrochloric acid soluble calcium.This implies that the plants that are relatively sensitive to drought hold more calcium ion,while the drought-resistance plants hold more calcium oxalate.Thus,the plant calcium components are in close relation to plant drought-resistance,and of important significance in plant physiology of the desert.

Human Causes of Aeolian Desertification in Northern China

Aeolian desertification has rapidly developed in the past 50 years in Northern China,covered an area of 0.386 million km2 by 2000,affected nearly 170 million population,and caused the direct and indirect economic loss of about $6.75(U.S.dollar) billion per year.Here we present several lines of evidence to demonstrate that human activities guided by policy shifts have been a major force to drive aeolian desertification via changes in land-use patterns and intensity.It is suggested that the desertification can be curbed or even reversed by adopting prevention and control measures with ecologically sound land-use practices in China.

Effects of sand burial on dune plants: a review

Burial of different growth stages of plants （e.g., adult plants, seedlings and seeds） is frequent in dune ecosystems. The soil micro-environment, which differs from surface conditions, influences the survival and growth of dune plants. To sum up knowledge about the survival mechanisms of plants under sand burial and to promote practical rehabilitation of dune vegetation, we reviewed relevant published literature and concluded that： （1） Focus in recent years has been on impacts of sand burial on seed germination and seedling emergence. Generally, shallow burial increased seed germination and seedling emergence, but deeper burial was negative. Buried at the same depth, large seeds showed higher germination and seedling emergence rates, attributed to larger energy reserves. （2） Survival, growth and reproduction rates of dune plants show plasticity in response to sand burial. Long-term deep burial is fatal because it creates a physical barrier which overcomes the vertical growth of plants, reduces photosynthetic leaf area, and limits oxygen availability to roots. Modest burial, on the other hand, is advantageous for growth and reproduction of many dune plants, due to protection from excessive temperature and drought. （3） There are few reports concerning effects of sand burial on plant physiology, but a limited number of studies indicate that partial burial increases water use efficiency, chlorophyll content, transpiration rate and net photosynthetic rates. The antioxidant protective enzyme system and osmolyte balance were reported to be involved in the mechanisms of dune plant resistance to burial.

Reproduction of Hedysarum scoparium (Fabaceae) in patched habitat is pollen limited,but not just pollinator limited

Pollen limitation of plant reproduction occurs across Angiosperms, particularly those in patched habitats We investigated the, relationship between pollen limitation and patch variables （patch size, visitation frequency） in the desert plant Hedysarum scoparium （Fabaceae）, which is an important xerophyte in the arid and semi-arid regions of Northwest China and can grow well as a pioneer plant in shifting sand dunes. We observed insect visitation to H. scoparium over two flowering seasons and estimated pollen limitation using fruit set and seed production. Our results indicate that fruit set and seed production increased significantly with pollen supplementation compared with open pollination. Hedysarum scoparium was pollinated by over 8 species of bees, with 88.4% of visits made by introduced honeybees （Apis mellifera）. Bee visitation varied significantly among the patches of habitats, but not associated with patch size of habitat. In general, pollen limitation occurred more strongly during fruit set than during seed production. The patches that received higher rates of pollinator visits were less pollen limited for fruit set. Pollen limitation for seed production, however, was not associated with pollinator visitation frequency. We conclude that pollen limitation in H. scoparium was caused by more than one reason, not just pollinator visits.

Effects of grazing and climate change on sandy grassland ecosystems in Inner Mongolia

To understand the effects of grazing activities and climate change on sandy grassland ecosystems in northem China, a livestock field grazing and enclosure experiment was conducted from 1992 to 2006 in Horqin Sand Land, Inner Mongolia. The results showed that sustained heavy grazing resulted in serious degradation of the vegetation; moderate grazing can maintain vegetation stabilization; and light grazing can promote rapid restoration of degraded vegetation. The livestock productivity was the highest in the moderate grazing grassland, and sustained heavy grazing resulted in rapid decrease of the livestock productivity. Heavy grazing can cause a retrogressive succession of grassland vegetation, whereas moderate and light grazing may promote progressive succession of plant species. The effects of changing climate on succession processes were not significant in the short term; a warm-humid climate is favorable to restoration of degraded vegetation, whereas a sustained warm-drought climate may result in degradation of grassland vegetation. Heavy livestock grazing should be stopped for the sustainable use of grassland; the proper grazing intensity for sandy grassland is two to three sheep or sheep equivalents per hectare in Inner Mongolia.

Effects of soil nitrogen:phosphorus ratio on growth rate of Artemisia ordosica seedlings

To address how the ratios of nitrogen and phosphorus (N:P ratios) in soil affect plant growth, we performed a two-factor (soil available N:P ratios and plant density) randomized block pot experiment to examine the relationships between soil N:P ratios, and the N:P ratios and growth rate of Artemisia ordosica seedlings. Under moderate water stress and adequate nutrient status, both soil N:P and plant density influenced the N:P ratios and growth rates of A. ordosica. With the increase of soil N:P ratios, the growth rates of A. ordosica seedlings decreased significantly. With the increase of soil N:P ratios, N:P ratios in A. ordosica seedlings increased significantly. While the nitrogen concentrations in the plant increased slightly, the phosphorus concentrations significantly decreased. With the increase of plant density, the shoot N:P ratios and growth rates significantly decreased, which resulted from soil N:P ratios. Thus, soil N:P ratios influenced the N:P ratios in A. ordosica seedlings, and hence, influenced its growth. Our results suggest that, under adequate nutrient environment, soil N:P ratios can be a limiting factor for plant growth.

Effects of soil nitrate:ammonium ratio on plant carbon:nitrogen ratio and growth rate of Artemisia sphaerocephala seedlings

Can soil nitrate： ammonium ratios influence plant carbon： nitrogen ratios of the early succession plant？ Can plant carbon： nitrogen ratios limit the plant growth in early succession？ To address these two questions, we performed a two-factor （soil nitrate： ammonium ratio and plant density） randomized block design and a uniform-precision rotatable central composite design pot experiments to examine the relationships between soil nitrate： ammonium ratios, the carbon： nitrogen ratios and growth rate of Artemisia sphaerocephala seedlings. Under adequate nutrient status, both soil nitrate： ammonium ratios and plant density influenced the carbon： nitrogen ratios and growth rate of A. sphaerocephala seedlings. Under the lower soil nitrate： ammonium ratios, with the increase of soil nitrate： ammonium ratios, the growth rates of plant height and shoot biomass of A. sphaerocephala seedlings decreased significantly; with the increase of plant carbon： nitrogen ratios, the growth rates of shoot biomass of A. sphaerocephala seedlings decreased significantly. Soil nitrate： ammonium ratios affected the carbon： nitrogen ratios of A. sphaerocephala seedlings by plant nitrogen but not by plant carbon. Thus, soil nitrate： ammonium ratios influenced the carbon： nitrogen ratios of A. sphaerocephala seedlings, and hence influenced its growth rates. Our results suggest that under adequate nutrient environment, soil nitrate： ammonium ratios can be a limiting factor for the growth of the early succession plant.

Effects of sand burial on growth and antioxidant enzyme activities of wheat(Triticum aestivum L.) in northern China

Growth and physiological responses of wheat to sand burial were studied in Horqin Sandy Land, to determine the impact on productivity and survival as well as antioxidant enzymes responses. This study consisted of one control （no sand） and four sand burial treatments： 25%, 50%, 75% and 100% of seedling height, respectively. Minor burial （25%） had no effect on wheat growth and survival; deep burial （100%） was fatal, and the others had an intermediate effect. Thus, the survival limit to sand burial was equal to seedling height. Sand burial mainly decreased shoot biomass and crop yield, but had small effects on belowground biomass. Superoxide dismutase （SOD） activity increased with time after burial in all treatments with surviving plants. Peroxidase （POD） activity increased after six days under burial, and catalase （CAT） activity de- creased after burial, but recovered after 12 days. The concentration of malondialdehyde （MDA）, a marker for oxidative stress, was low on the sixth day, but increased thereafter with burial depth. Thus, sand burial 〉25% should be avoided due to growth rate reduction leading to reduced crop yield, and even 25% burial showed physiological indicators of stress.

Effects of sand burial on survival and growth of Artemisia halodendron and its physiological response

There is a great deal of literature on the effects of sand burial upon the survival and growth of desert plants, but the physiological adaption mechanisms of desert plants to sand burial have as yet rarely been studied. Artemisia halodendron is widely distributed in the semi-arid deserts of China and is a dominant species in semi-moving dune vegetation. The growth and physiological properties ofA. halodendron seedlings under different sand burial depths were studied in 2010 and 2011 in the Horqin Sand Land, Inner Mongolia, to better understand the ability and physiological mechanism by which desert plants withstand sand burial. The results showed that A. halodendron as a prammophyte species had a stronger ability to withstand sand burial compared to non-prammophytes, with some plants still surviving even if buried to a depth reaching 225% of seedling height. Although seedling growth was inhibited significantly once the depth of sand burial reached 50% of the seedling height, seedling survival did not decrease significantly until the burial depth exceeded 100% of the seedling height. Sand burial did not result in significant water stress or MDA （Malondialdehyde） accumulation in the seedlings, but membrane permeability increased significantly when the burial depth exceeded 100% of the seedling height. After being subjected to sand burial stress, POD （Peroxidase） activity and proline content increased significantly, but SOD （Superoxide Dismutase） and POD activities and soluble sugar content did not. The primary mechanism resulting in in- creased mortality and growth inhibition were that cell membranes were damaged and photosynthetic area decreased when subjected to the severe stress of sand burial, while proline and POD played key roles in osmotic adjustment and protecting cell membranes from damage, respectively.

Seasonal change mediates the shift between resource and pollen limitation in Hedysarum scoparium(Fabaceae)

Theory suggests that with sufficient environmental variation, pollen limitation might be observed at some places or times, and resource limitation at others, but there are no empirical data about the effect of seasonal change on the variation of pollen limitation and resource limitation within a flowering season. In this study, we examined pollen and resource limi- tation by comparing fruit set and seed production in natural- and hand-pollinated Hedysarum scoparium flowers in the middle reaches of the Hexi Corridor region, China, in 2010. We also described a role for the first substantial autumn rainfall in mediating a shift between pollen and resource limitation in H. scoparium, but did not analyze this experimentally Our results indicated that H. scoparium was resource limited at peak flowering during the summer, and was pollen limited at peak flowering during the autumn. The seasonal change （summer to autumn） mediated the shift between pollen and resource limitations in H. scoparium. The shift timing depended on the date of the first autumn rainfall in 2010. Changes in the first substantial rainfall in autumn may affect fruiting of H. scoparium, thus affecting population persistence of this species and development/structure of the local ecosystem if such conditions persist.