The Qinghai-Xizang Plateau and its adjacent highlands host a rich plant diversity, and understanding on how plant species reproduce successfully in those high mountain regions would be of great help for us to discover the evolution and adaptability of plants to the typical alpine environments. Here, we reviewed the progresses of plant reproductive ecology in the Qinghai-Xizang Plateau and its adjacent highlands, with emphases on resource allocation, reproductive mode, evolution of floral traits, etc. We also summarized the impacts of global climate changes on plant reproduction, and the applications of new technologies and methods in this research field to explore the potential mechanisms of plant evolution and adaptation. In the stressful environments of high mountain areas, with the increase of altitude, the reproductive allocation usually shows a trend of increase, and the proportion of resources invested in male function also increases. However, the specific pattern of resource allocation strongly depends on the mating system, individual size, life history, genetic variation and resource availability. In the context of resource and pollination limitation, plants will weight different reproductive modes. Clonal reproduction and self-pollination are conductive to reproductive assurance in populations with pollinator scarcity, while sexual reproduction and cross-pollination can improve seed quality and genetic diversity of offspring, which could benefit population survival in the potential climate flunctations. Therefore, the combinations of different reproductive modes and the generalized pollination networks may be the good choices to adapt to the stressful environments of highlands. The evolution of floral traits might be mainly driven by the selective pressure of local pollinators, but more and more attentions have been paid to the effects of exotic pollinators, herbivores, nectar robbers and abiotic environments such as temperature, precipitation and ultraviolet radiation on floral traits. In recent years, the Qinghai-Xizang Plateau has become one of research hotspots because of its fragility and sensitivity to climate change. Global climate change, characterized by global warming and increased nitrogen deposition, is directly or indirectly affecting the reproductive characteristics of plant species in the region. Climate change may cause shifts of phenology and distribution area of plants and pollinators, which leads to temporal decoupling and spatial mismatch between them. Plants can respond to climate change by adjusting floral traits (floral display, corolla structure, quantity and quality of reward), which may result in changes of type, amount and foraging behaviors of pollinators and affect reproductive success of plants at last. The applications of new technologies and methods could advance our knowledge on plant reproductive ecology. 3D printed flowers could control floral traits precisely and could be employed in examining the effects of subtle changes of floral traits on pollinator behaviors, which would be useful in understanding pollinator mediated selection on floral traits in combination with the uses of commercial pollinators. Importantly, with the development of high-throughput sequencing, the underlying mechanisms of important topics in plant reproductive ecology could be discovered more easily than before, especially in the field of floral evolution. As a conclusion, key issues that need additional attention in the current research as well as the fields worthy of further study are highlighted.

Aims Nitrogen (N) deposition has made great impacts on the structure and function of terrestrial ecosystems in recent decades, and N is known as the main limiting element of plant growth and primary productivity in the northern grassland of China. Species diversity and functional diversity have been widely recognized as key indicators of the mechanisms of biodiversity maintenance for ecosystem functions. However, the effect of long-term N addition on the relationship between species diversity and functional diversity, and the corresponding impacts on primary productivity have rarely been studied; particularly the underpinning mechanisms remain unclear. Our objective is to examine the effects and pathways of long-term N addition on species diversity, functional diversity and primary productivity in a typical steppe.
Methods A manipulative N addition experiment located in a typical steppe of the Nei Mongol grassland has been conducted for 18 years. The experimental design included seven levels of N addition rate (i.e., control, 0, 1.75, 5.25, 10.50, 17.50, 28.00 g·m^-2·a^-1) with nine replicates for each treatment. The plant functional traits of dominant species, species richness and composition, and aboveground net primary productivity (ANPP) were determined. In addition, species diversity, functional attribute diversity and community-weighted mean traits were calculated.
Important findings 1) N addition significantly reduced species richness and Shannon-Wiener index, but had little effect on functional diversity, including functional attribute diversity and community-weighted mean traits. 2) Structural equation model analyses showed that functional diversity was mainly affected by species richness, whereas the decrease in species diversity didn’t lead to the decrease in functional diversity. The community- weighted mean traits did not change under N addition, which was mainly due to the shift in functional group composition, that is, the proportion of perennial rhizome grass in plant community increased greatly under N addition. 3) The effect of N addition on ANPP was through direct pathways of species richness and shift in functional group composition, and further via an indirect pathway of community-weighted mean traits. Particularly, the community-weighted mean traits were the most important factors and explained 48% of total variation in ANPP, implying that the primary productivity is mainly determined by the biomass and functional traits of dominant species in the plant community. Our results well supported the mass-ratio hypothesis.

Aims Shrub encroachment is a critical ecological problem in arid and semi-arid ecosystems worldwide. The effects of shrub encroachment on ecosystem structure and function of grasslands are complicated and need to be explored in future studies. Our objective is to examine the effects and pathways of shrub encroachment on ecosystem structure and function in a typical steppe of the Inner Mongolia grassland.
Methods Three grassland sites with different degrees of shrub encroachment (i.e. light, moderate, heavy) were selected in the Xilingol Nei Mongol, of which Caragana microphylla was the dominant shrub. Species richness and composition, aboveground net primary productivity (ANPP), soil property, and plant functional traits of dominant species were determined in this study. In addition, species diversity, functional attribute diversity, community-weighted mean traits, and vegetation leaf and soil carbon and nitrogen pools were further calculated.
Important findings 1) The species richness, functional attribute diversity and community-weighted mean traits differed significantly among three grassland sites, and species diversity and functional diversity were relatively higher in the moderate shrub-encroachment site, indicating moderate shrub-encroachment favors biodiversity maintenance. 2) The aboveground net primary productivity of heavy shrub-encroachment grassland was significantly higher than those of light and moderate shrub-encroachment grasslands, which was mainly due to a shift in functional group composition, that is, the proportion of annuals and biennials to perennial grasses and forbs increased greatly with intensifying shrub encroachment. The vegetation leaf and soil carbon and nitrogen pools differed little among three sites. 3) Shrub encroachment did not directly affect ecosystem function, including ANPP, vegetation and soil nutrient pools, but it indirectly affected them through pathways of the shift in functional group composition and changes in soil property and functional diversity. Particularly, the shift in functional group composition and intensified soil drought and basification was separately important biotic and abiotic factors for variations in ecosystem function.

Aims Tsoongiodendron odorum is an ancient relic species belonging to the family Magnoliaceae, but it is labelled endangered plant with extremely small populations and facing serious threats to its wild survival now. Using Ecological Niche Modelling (ENM) to hindcast historical changes in its distribution during the Last Glacial Maximum (LGM), this study aims to explore the impact of climate change on the distribution of T. odorum, and to evaluate the relationship between species distribution and environmental variables. The results of this study could contribute to the conservation of T. odorum in the context of global warming.
Methods Based on 96 modern geographical distribution records and 8 bioclimatic variables, we simulated the potential distribution of T. odorum during the LGM, Mid-Holocene, present and future (period of 2061-2080 in the Representative Concentration Pathway 8.5 climate scenario) with MaxEnt model. The changes in species distribution through time were analyzed by SDM toolbox, while the importance of bioclimatic variables was evaluated by percent contribution, permutation importance and Jackknife test.
Important findings (1) The highly suitable region for T. odorum was Nanling region, and this area might be the refuge where T. odorum survived in situ during the LGM because only slightly southward retreat in distribution was detected in this region during the LGM. (2) In the two warming climate scenarios (Mid-Holocene and future), the area of the suitable region was reduced, and the decrease of future distribution is greater than that during the Mid-Holocene, which suggests that warming climate might have a negative impact on the distribution of T. odorum. (3) Overall the stability of distribution range of T. odorum in each period indicates the climate adaptation of this species. Human activity or self-breeding problem was likely the significant cause leading to endangered condition. Guangdong and Guangxi should be regarded as priority conservation areas as shown by our results.

Aims Drought is a limiting factor for plant growth in southern karst areas. Climate change may affect the amount and distribution pattern of precipitation in these areas. It is important to understand the stoichiometric characteristics of soil and plants and how they respond to increasing precipitation in karst areas.
Methods In Jianshui karst areas in southern Yunnan, a water addition experiment was conducted since April 2017 and the concentrations of carbon (C), hydrogen (H), nitrogen (N), phosphorus (P), sulfur (S), potassium (K), calcium (Ca), magnesium (Mg), aluminum (Al), sodium (Na), iron (Fe), manganese (Mn), zinc (Zn), copper (Cu) in the soils and leaves of two dominant shrubs (Bauhinia brachycarpa and Carissa spinarum) were measured in the dry season (April) and rainy season (August) in 2018.
Important findings Water addition affected the content of C, N and Na in the soil. Compared with the dry season, the concentrations of Na and S in the soil significantly decreased in the rainy season. The remaining soil elements did not show any significant differences between treatments and seasons. With the increase of soil moisture content, the concentrations of K decreased while Ca in both plant species increased. These results also indicated that soil moisture changes could significantly affect plant ecological traits. With soil moisture changes, the stabilities of leaf elements were related to their contents. The closer the leaf element contents are to the corresponding maximum or minimum values, the smaller are the coefficients of variation. And the variation coefficients of P, S and Mg with the concentrations close to 1 mg·g^-1were the highest. Under the changes of soil moisture conditions, the stability of C, N, P and other major elements in C. spinarum was significantly higher than that B. brachycarpa. Changes in soil water content, which was caused by both rainfall changes and water addition, had different effects on different the contents of different elements in both soil and plants. These results may shed light on the restoration of soil and plants in karst regions.

Aims The agro-pastoral ecotone is considered as fragile ecosystems which are strongly affected by agriculture and animal husbandry. The saline-alkali grassland is a unique grassland type in the agro-pastoral ecotone. A large amount of fertilizers are used to increase productivity in this area, which also promotes the emission of reactive nitrogen (N) gases and leads to the changes in soil carbon and N cycles. Mowing is a primary management practice in the agro-pastoral grassland in northern China. In order to explore the impact of N addition and mowing on carbon dynamic in this saline-alkali grassland located in the agro-pastoral ecotone, we determined the response of soil respiration to N addition and mowing.
Methods This study area is located in Youyu County, an agro-pastoral grassland ecosystem in northern China. The field experiment was set up in May, 2017. The treatments included: control (without mowing and mowing), addition of urea, addition of slow release urea, addition of urea + mowing, addition of slow release urea + mowing. Each treatment included 6 replicates. Therefore, there were totally 36 plots in this experiment. Soil respiration rate, soil temperature, soil moisture content, microbial biomass, inorganic N content, above-ground and below-ground biomass were measured under different treatments, and the cumulative carbon emissions and CO₂ fluxes were calculated.
Important findings Our results showed that: (1) Short-term (2017-2018) N addition significantly increased soil respiration rates and soil cumulative carbon emissions. Meanwhile, soil respiration rates and cumulative carbon emissions were significantly higher under urea treatment than those under slow release urea addition. (2) Mowing significantly reduced soil respiration rates and cumulative carbon emissions. (3) The interaction of short-term N addition and mowing had no significant effect on soil respiration rate. Therefore, short-term N addition can promote soil carbon release from the saline-alkali grassland in the agro-pastoral ecotone of northern China. Mowing can reduce soil respiration and decrease cumulative of carbon emissions. This may be because that mowing reduced the input of litter and further reduced soil substrate for microbes, which led to a decrease in soil microbial activity. However, long-term effect of N addition and mowing on soil carbon dynamics in saline-alkaline grasslands in the agro-pastoral ecotone still needs to be further explored.

Aims Soil respiration is an important indicator for evaluation of ecosystem health in the grazing grasslands of arid regions, and thus can be used to assess dynamics of ecosystem functioning during the restoration of degraded grasslands from enduring intensive grazing.
Methods This study was carried out in a Nei Mongol desert grassland with four grazing intensity treatments, i.e., control, light, moderate, and heavy grazing intensity designated as CK, LG, MG, and HG, respectively. Our objectives of this study were to explore the responses of soil respiration in these treatments with additional nitrogen (N) and water (W) addition. The plant community was dominated by a grass species, Stipa breviflora.
Important findings Our results showed that: (1) previous grazing intensity had significant impacts on soil respiration in 2016 and 2017, but not in 2015. Grazing increased soil respiration. Moreover, both nitrogen and water addition significantly enhanced soil respiration in MG plots, while only combined addition of nitrogen and water significantly increased soil respiration in HG plots. (2) Neither grazing intensity nor addition of nitrogen and water changed the seasonal dynamics of growing season soil respiration in this desert grassland. Soil respiration showed a single-peak curve model, and the peak occurred in July with both rain and heat. (3) The effects of nitrogen and water addition varied in different growing seasons. Nitrogen addition had no significant effects in the first two years (2015 and 2016), while showed significant effects in the third year (2017). Water addition had significant effects in years with normal precipitation (2015 and 2017), while had insignificant effect in the year with high precipitation (2016). Combined addition of nitrogen and water showed stronger effects than only addition of water in CK, LG, and HG plots, indicating that the synergistic effects of nitrogen and water addition on soil respiration. (4) The sensitivity of soil respiration to soil temperature at 10 cm depth (i.e., the Q₁₀ value) ranged between 1.13 and 2.41, with an average value of 1.71. Without addition of nitrogen and water, Q₁₀ values in grazing plots were all lower than in CK plots, with the lowest value occurring in HG plots. With the addition of water and combined addition of water and nitrogen, the Q₁₀ value increased significantly by 100%. Taken together, our results indicated that soil moisture was the leading environmental factor affecting soil respiration in this desert grassland, while nitrogen played an effective role only after the minimum requirement of water availability was met. Results from this study will provide important helpful information for the restoration and rational utilization of the degraded desert steppe.