Hong-Yu Guo

Distribution pattern of Caragana species under the influence of climate gradient in the Inner Mongolia region, China

There is a strong climate gradient in the Inner Mongolia region, China, with solar radiation and air temperature increasing but precipitation decreasing gradually from the northeast to the southwest. Sixteen Caragana species exist in the Inner Mongolia region. These Caragana species exhibit a distribution pattern across moisture zones and form a geographical replacement series. In order to examine the mechanisms responsible for Caragana species distribution pattern, we selected 12 Caragana species that exhibit a distinct distribution pattern across multiple moisture zones in the Inner Mongolia region, and determined the relationships between the leaf ecological and physiological traits of these Caragana species and the aridity index and solar radiation. Along with the climatic drought gradient and the solar radiation intensification from the northeast to the southwest, leaf ecological characteristics of Caragana species changed drastically, i.e. the leaf shape gradually turned from flat into tegular or tubbish; the leaf hair became denser, longer and lighter in color; the leaf area, leaf biomass and specific leaf area (SLA) decreased significantly; the leaf thickness and the ratio of leaf thickness to leaf area increased significantly; and the leaf chlorophyll content decreased significantly. As the climatic drought stress increased, osmotic potentials of the main osmotic adjustment substances and the cytoplasmic ion concentration of Caragana species increased significantly. Meanwhile, the total and free water contents and water potential of leaves decreased significantly; the ratio of bound to free water increased significantly; the stomatal conductance and transpiration rate reduced significantly; and the water use efficiency (WUE) increased significantly. In addition, with the intensification of climatic drought stress, peroxidase (POD) and superoxide dismutase (SOD) activities in leaves increased significantly. As a result, the malondialdehyde (MDA) content increased while the oxygen free radical content decreased. Our results showed that most of the leaf ecological and physiological traits of the 12 Caragana species varied in accordance with the climatic drought gradient in the Inner Mongolia region, which reflected the adaptation of the Caragana species to the local climate conditions. With relatively more active metabolism and faster growth, the Caragana species in the northeast had strong competitive abilities; on the other hand, with stronger resistance to climatic drought stress, the Caragana species in the southwest could survive in harsh environments. Based on our results, we concluded that both the environmental gradients and the adaptive responses of Caragana species to their environments played important roles in the formation of the Caragana species distribution pattern across the Inner Mongolia region.

Changes in spatial patterns of Caragana stenophylla along a climatic drought gradient on the Inner Mongolian Plateau.

Few studies have investigated the influence of water availability on plant population spatial patterns. We studied changes in the spatial patterns of Caragana stenophylla along a climatic drought gradient within the Inner Mongolian Plateau, China. We examined spatial patterns, seed density, “nurse effects” of shrubs on seedlings, transpiration rates and water use efficiency (WUE) of C. stenophylla across semi-arid, arid, and intensively arid zones. Our results showed that patches of C. stenophylla populations shifted from a random to a clumped spatial pattern towards drier environments. Seed density and seedling survival rate of C. stenophylla decreased from the semi-arid zone to the intensively arid zone. Across the three zones, there were more C. stenophylla seeds and seedlings underneath shrub canopies than outside shrub canopies; and in the intensively arid zone, there were almost no seeds or seedlings outside shrub canopies. Transpiration rates of outer-canopy leaves and WUE of both outer-canopy and inner-canopy leaves increased from the semi-arid zone to the intensively arid zone. In the intensively arid zone, transpiration rates and WUE of inner-canopy leaves were significantly lower and higher, respectively, than those of outer-canopy leaves. We conclude that, as drought stress increased, seed density decreased, seed proportions inside shrubs increased, and “nurse effects” of shrubs on seedlings became more important. These factors, combined with water-saving characteristics associated with clumped spatial patterns, are likely driving the changes in C. stenophylla spatial patterns.

Acclimation of Photosynthetic Traits of Caragana Species to Desert Environment in Inner Mongolian Plateau

Caragana species are commonly distributed in desert regions. We hypothesized that the photosynthetic traits of desert Caragana species could acclimated well to the desert environments, which are characterized by high temperature, intense radiation, and low moisture. To test our hypothesis, we studied the responses of photosynthesis to light, temperature and humidity, and fluorescence parameters of four Caragana species from desert region and one Caragana species from steppe region in Inner Mongolian Plateau. The desert Caragana species had higher light compensation point, light saturation point, optimum temperature for photosynthesis, and lower optimum relative air humidity for photosynthesis than the steppe Caragana species, and their net photosynthesis rates (Pn) were higher under high temperature, intense sunlight, and low humidity. The nonphotochemical quenching efficiency (NPQ) and minimal fluorescence (Fo) of the desert Caragana species were higher, and the values of maximum fluorescence (Fm), maximum quantum yield (Fv/Fm), relative quantum yield (ΦPSII), and photochemical quenching coefficient (qP) were lower, except for C. korshinskii. The electron transport rate (ETR) of the desert Caragana species was greater or similar to that of C. microphylla. In summary, the photosynthetic traits of the desert Caragana species acclimated well to desert environment, although such acclimation might be still not sufficient. The order of photosynthetic acclimation of the four species to desert environment is: C. korshinskii > C. stenophylla > C. tibetica > C. roborovskyi.

Effects of grazing intensity on seed production of Caragana stenophylla along a climatic aridity gradient in the Inner Mongolia Steppe, China

As the increases of climatic aridity and grazing intensity, shrubs play an increasingly important role in grassland ecosystem in arid and semi-arid regions, and its abundance also generally increases. However, the effects of climatic aridity and grazing intensity on sexual reproduction of shrubs in grassland remain largely unclear. In order to understand the effects of grazing intensity and climatic drought stress, and their interaction on seed production of shrub species, we examined the seed number, seed weight and seed yield of Caragana stenophylla under three grazing intensities (fenced, mild grazing and severe grazing) across a climatic aridity gradient (semi-arid, arid, very arid and intensively arid zones) in the Inner Mongolia Steppe, northern China during 2012–2013. The seed number, seed weight and seed yield gradually increased from the semi-arid to the very arid zones, but decreased from the very arid to the intensively arid zones in fenced plots. The seed number and seed yield decreased from the semi-arid to the intensively arid zones in mild and severe grazing treatment plots, therefore, grazing enhanced the suppression effect of climatic aridity on seed production of C. stenophylla. The seed number and seed yield gradually decreased as grazing intensity increased. The seed weight was highest in severe grazing plots, followed by the mild grazing plots and then the fenced plots. Precipitation varied interannually during the study period. We observed that the seed number, seed weight and seed yield were lower in the low precipitation year (2013) than in the high precipitation year (2012). As climatic drought stress increased, the negative effects of grazing on seed production of C. stenophylla also gradually increased. Our results indicated that climatic drought stress may contribute to the encroachment of C. stenophylla shrub in arid zones by promoting its seed production. However, grazing had negative effects on sexual reproduction of C. stenophylla, and the combined effects of drought stress and grazing seriously suppressed sexual reproduction of C. stenophylla in the intensively arid zone.

Alterations in flowering strategies and sexual allocation of Caragana stenophylla along a climatic aridity gradient

Plant can alter reproductive strategies for adaptation to different environments. However, alterations in flowering strategies and sexual allocation for the same species growing in different environments still remain unclear. We examined the sexual reproduction parameters of Caragana stenophylla across four climatic zones from semi-arid, arid, very arid, to intensively arid zones in the Inner Mongolia Steppe, China. Under the relatively favorable climatic conditions of semi-arid zone, C. stenophylla took a K-strategy for flowering (fewer but bigger flowers, and higher seed set). In contrast, under the harsher climatic conditions of intensively arid zone, C. stenophylla took an r-strategy for flowering (more but smaller flowers, and lower seed set). In arid and very arid zones, C. stenophylla exhibited intermediate flowering strategies between K- and r-strategies. In semi-arid, arid and very arid zones, sexual allocation and sexual allocation efficiency (SAE) of C. stenophylla were high, and the population recruitment might be mainly through sexual reproduction; in intensively arid zone, however, sexual allocation and SAE were very low, seed production was very limited, and clonal reproduction might compensate for the decrease in sexual reproduction. Our results suggested that C. stenophylla adapted to the climatic aridity gradient by alterations in flowering strategies and reproductive allocation.

Effects of Grazing on Population Growth Characteristics of Caragana stenophylla Along a Climatic Aridity Gradient

ABSTRACT Shrubs are important plant species in grassland ecosystems worldwide, and their density and cover have been gradually increasing globally. However, the interaction effect of grazing and aridity on population recruitment and population growth of shrub species in grasslands has not been examined explicitly. We examined sapling establishment, sexual recruitment, population mortality, and population growth of Caragana stenophylla along a climatic aridity gradient and a grazing intensity gradient in the Inner Mongolia Steppe, using manipulative field experiments. Sapling establishment, sapling height, and sexual recruitment of C. stenophylla decreasedas climatic aridity and grazing intensity increased. The negative effects of grazing on sapling establishment and sexual recruitment gradually increased as climatic aridity increased. The effect of climatic aridity and grazing on population mortality was influenced by sexual recruitment. In the combined treatments of climatic aridity and grazing, population mortality was relatively high when sexual recruitment was relatively high, while population mortality increased as climatic aridity and grazing increased when sexual recruitment was relatively low. C. stenophylla population increased under relatively low drought stress and mild grazing but declined under strong drought stress and/or severe grazing. Our results suggested that to maintain viable Caragana populations, appropriate grazing policies must be made according to climate aridity gradient.

What drives the shift between sexual and clonal reproduction of Caragana stenophylla along a climatic aridity gradient

BackgroundThe reasons that clonal plants shift between sexual and clonal reproduction have persisted as a knowledge gap in ecological literature. We hypothesized that clonal plants’ shifts between sexual and clonal reproduction in different environments are driven by the relative costs of sexual and clonal reproduction. Moreover, we hypothesized plants prioritize sexual reproduction over clonal reproduction. To test these hypotheses, we determined the costs of sexual and clonal reproduction, and proportions of sexual and clonal reproduction of Caragana stenophylla along a climatic aridity gradient (semi-arid, arid, very arid and intensively arid zones) in the Inner Mongolia Steppe using several complementary field experiments.ResultsThe cost of sexual reproduction increased while the cost of clonal reproduction decreased as climatic drought stress increased from the semi-arid to the intensively arid zones. The changes in the costs of these reproductive modes drove a shift in the reproductive mode of C. stenophylla from more sexual reproduction in the semi-arid zone to more clonal propagation in the intensively arid zone. However, because of the evolutionary advantages of sexual reproduction, sexual reproduction still held priority over clonal production in C. stenophylla, with the priority of sexual reproduction gradually increasing from the semi-arid to the intensively arid zones.ConclusionsOur study suggested that sexual reproduction has relatively high priority in propagation of C. stenophylla. However, if the costs of sexual reproduction are too high, C. stenophylla likely chooses clonal reproduction, and the ratio between sexual and clonal reproduction could be mediated by reproductive cost. These reproductive strategies reflect optimal resource utilization, and allow the persistence of both reproductive modes across stressful conditions depending on their evolutionary advantages.

Grazing limits natural biological controls of woody encroachment in Inner Mongolia Steppe

ABSTRACT Woody encroachment in grasslands has become increasingly problematic globally. Grazing by domestic animals can facilitate woody encroachment by reducing competition from herbaceous plants and fire frequency. Herbivorous insects and parasitic plants can each exert forces that result in the natural biological control of encroaching woody plants through reducing seeding of their host woody plants. However, the interplay of grazing and dynamics of herbivorous insects or parasitic plants, and its effects on the potential biological control of woody encroachment in grasslands remains unclear. We investigated the flower and pod damage by herbivorous insects, and the infection rates of a parasitic plant on the shrub Caragana microphylla, which is currently encroaching in Inner Mongolia Steppe, under different grazing management treatments (33-year non-grazed, 7-year non-grazed, currently grazed). Our results showed that Caragana biomass was highest at the currently grazed site, and lowest at the 33-year non-grazed site. Herbaceous plant biomass followed the opposite pattern, suggesting that grazing is indeed facilitating the encroachment of Caragana plants in Inner Mongolia Steppe. Grazing also reduced the abundance of herbivorous insects per Caragana flower, numbers of flowers and pods damaged by insect herbivores, and the infection rates of the parasitic plant on Caragana plants. Our results suggest that grazing may facilitate woody encroachment in grasslands not only through canonical mechanisms (e.g. competitive release via feeding on grasses, reductions in fires, etc.), but also by limiting natural biological controls of woody plants (herbivorous insects and parasitic plants). Thus, management efforts must focus on preventing overgrazing to better protect grassland ecosystems from woody encroachment. Summary: Grazing can negatively affect herbivorous insects and parasitic plants on woody plants, and thus limit the effectiveness of their natural biological control effects on woody encroachment.

Shrubs facilitate recruitment of Caragana stenophylla Pojark: microhabitat amelioration and protection against herbivory

Key messageMature Caragana stenophylla shrubs facilitated intraspecific sapling establishment by two mechanisms: microhabitat amelioration and protection against herbivory. Facilitation was mediated by climate, grazing, and sapling age.ContextPre-existing shrubs could facilitate sapling establishment of woody plants; however, how these facilitation vary across abiotic and biotic stress gradients and the underlying mechanisms remain unclear.AimsThe aim of this study is understanding the facilitation of shrub on sapling establishment and how the two underlying mechanisms, microhabitat amelioration and protection against herbivory, vary across climatic aridity gradients, grazing gradients, and sapling age.MethodsWe conducted field sowing experiments to examine the facilitation of mature Caragana stenophylla Pojark on intraspecific sapling establishment.ResultsFacilitation of C. stenophylla on sapling survival increased as drought stress, grazing intensity, and sapling age increased. Microhabitat amelioration increased as drought stress and sapling age increased. Similarly, protection against herbivory increased as drought stress, grazing intensity, and sapling age increased. Relative importance of microhabitat amelioration increased as drought stress increased, and relative importance of protection against herbivory increased as grazing intensity and sapling age increased.ConclusionFacilitation of shrub on sapling establishment involves both microhabitat amelioration and protection against herbivory. Facilitation, the two mechanisms, and relative importance between the two mechanisms would all be affected by climatic aridity, grazing intensity, and sapling age. Shrub establishment has a positive feedback effect.