Trunk volume estimation of irregular shaped Populus euphratica riparian forest using TLS point cloud data and multivariate prediction models

Background: Trunk volume(Vt) is an essential parameter for estimating forest stand volume, biomass, and carbon sequestration potential. As the dominant tree species in desert riparian forests, Euphrates poplar(Populus euphratica) has a high proportion of irregularly shaped tree trunks along the Tarim River, NW China, where the habitat is very fragile owing to long-term water stress. This causes uncertainty in estimation accuracy as well as technical challenges for forest surveys. Our study aimed to acquire P. euphratica Vtusing terrestrial laser scanning(TLS) and to establish a species-specific Vtprediction model.Methods: A total of 240 individual trees were measured by TLS multiple-station in 12 sampling plots in three sections along the lower reaches of the Tarim River. Vtwas calculated by a definite integration method using trunk diameters(Di) at every 0.1-m tree height obtained from TLS, and all data were split randomly into two sets:70% of data were used to estimate the model parameter calibration, and the remaining 30% were used for model validation. Sixteen widely used candidate tree Vtestimation models were fitted to the TLS-measured Vtand tree structural parameter data, including tree height(H), diameter at breast height(DBH), and basal diameter(BD). All model performances were evaluated and compared by the statistical parameters of determination coefficient(R^(2)),root mean square error(RMSE), Bayesian information criterion(BIC), mean prediction error(ME), mean absolute error(MAE), and modeling efficiency(EF), and accordingly the best model was selected.Results: TLS point cloud reflection intensity(RI) has advantageous in the extraction of data from irregular tree trunk structures. The P. euphratica tree Vtvalues showed obvious differences at the same tree height(H). There was no significant correlation between Vtand H(R^(2)=0.11, P < 0.01), which reflected the irregularity of P. euphratica trunk shape in the study area. Among all the models, model(14): Vt=0.909DBH1.184H0.487BD0.836(R^(2)=0.97, RMSE=0.14) had the best prediction capability for irregularly shaped Vtwith the highest R^(2), BIC(-37.96), and EF(0.96), and produced a smaller ME(0.006) and MAE(1.177) compared to other models. The prediction accuracy was 93.18%.Conclusions: TLS point cloud RI has a potential for nondestructively measuring irregularly shaped trunk structures of P. euphratica and developed Vtprediction models. The multivariate models more effectively predicted Vtfor irregularly shaped trees compared to one-way and general volume models.

Structural Characteristics and Eco-adaptability of Heteromorphic Leaves of Populus euphratica

The microstructural and ultrastructural traits of three kinds of typical leaves of Populus euphratica Olive, including lanceolate, broad-ovate and dentate broad-ovate leaves, were studied by using electron microscope and optical microscope. The re- sults showed that with the leaves changing from lanceolate shape to dentate broad-ovate shape, their structure obviously tended to be xeromorph: developed palisade tissue, undeveloped spongy tissue, thick cutin layer and sunken stomas. The amount of mitochondria tended to be increased, and the shape of chloroplasts varied from regular spindle to irregular rotundity or oval. The leaves were cov- ered with wax without cilium, and the stomas on the upper and lower epidermis of the leaves opened unevenly. The stomas on the lower epidermis were deeper than those on the upper epidermis under the scanning electron microscope. The results implied that the structural characteristics of the diversiform-leaves of P. euphratica are related to its eco-adaptability.

Stand structure and height-diameter relationship of a degraded Populus euphratica forest in the lower reaches of the Tarim River, Northwest China

Understanding stand structure and height-diameter relationship of trees provides very useful information to establish appropriate countermeasures for sustainable management of endangered forests. Populus euphratica, a dominant tree species along the Tarim River watershed, plays an irreplaceable role in the sustainable development of regional ecology, economy and society. However, as the result of climate changes and human activities, the natural riparian ecosystems within the whole river basin were degraded enormously, particularly in the lower reaches of the river where about 320 km of the riparian forests were either highly degraded or dead. In this study, we presented one of the main criteria for the assessment of vitality of P. euphrafica forests by estimating the defoliation level, and analyzed forest structure and determined the height-diameter （height means the height of a tree and diameter means the diameter at breast height （DBH） of a tree） relationship of trees in different vitality classes （i.e. healthy, good, medium, senesced, dying, dead and fallen）. Trees classified as healthy and good ac- counted for approximately 40% of all sample trees, while slightly and highly degraded trees took up nearly 60% of total sample trees. The values of TH （tree height） and DBH ranged from 0-19 m and 0-125 cm, respectively. Trees more than 15 m in TH and 60 cm in DBH appeared sporadically. Trees in different vitality classes had different distribution patterns. Healthy trees were mainly composed more of relatively younger trees than of degraded tress. The height-diameter relationships differed greatly among tress in different vitality classes, with the coefficients ranging from 0.1653 to 0.6942. Correlation coefficients of TH and DBH in healthy and good trees were higher than those in trees of other vitality classes. The correlation between TH and DBH decreased with the decline of tree vitality. Our results suggested that it might be able to differentiate degraded P. euphratica trees from healthy trees by determining the height-diameter correlation coefficient, and the coefficient would be a new parameter for detecting degradation and assessing sustainable management of floodplain forests in arid regions. In addition, tree vitality should be taken into account to make an accurate height-diameter model for tree height prediction.

Gas exchange of Populus euphratica leaves in a riparian zone

Riparian vegetation belts in arid regions of Central Asia are endangered to lose their ecosystem services due to intensified land use.For the development of sustained land use,management knowledge of plant performance in relation to resource supply is needed.We estimated productivity related functional traits at the edges of the habitat of Populus euphratica Oliv.Specific leaf area （SLA） and carbon/nitrogen （C/N） ratio of P.euphratica leaves growing near a former river bank and close to moving sand dunes in the Ebinur Lake National Nature Reserve in Xinjiang,Northwest China （near Kazakhstan） were determined and daily courses of CO2 net assimilation （PN）,transpiration （E）,and stomatal conductance （gs） of two consecutive seasons were measured during July-August 2007 and June-July 2008.Groundwater level was high （1.5-2.5 m below ground） throughout the years and no flooding occurred at the two tree stands.SLA was slightly lower near the desert than at the former river bank and leaves contained less N in relation to C.Highest E and gs of P.euphratica were reached in the morning before noon on both stands and a second low maximum occurred in the afternoon despite of the unchanged high levels of air to leaf water vapor pressure deficit （ALVPD）.Decline of gs in P.euphratica was followed by decrease of E.Water use efficiency （WUE） of leaves near the desert were higher in the morning and the evening,in contrast to leaves from the former river bank that maintained an almost stable level throughout the day.High light compensation points and high light saturation levels of PN indicated the characteristics of leaves well-adapted to intensive irradiation at both stands.In general,leaves of P.euphratica decreased their gs beyond 20 Pa/kPa ALVPD in order to limit water losses.Decrease of E did not occur in both stands until 40 Pa/kPa ALVPD was reached.Full stomatal closure of P.euphratica was achieved at 60 Pa/kPa ALVPD in both stands.E through the leaf surface amounted up to 30％ of the highest E rates,indicating dependence on water recharge from the ground despite of obviously closed stomata.A distinct leaf surface temperature （Tleaf） threshold of around 30℃ also existed before stomata started to close.Generally,the differences in gas exchange between both stands were small,which led to the conclusion that micro-climatic constraints to E and photosynthesis were not the major factors for declining tree density with increasing distance from the river.

Isolation of protoplast from callus of Populus euphratica and H^+ fluxes across plasma membrane under NaCl stress

We used callus of Populus euphratica Olive to isolate protoplasts, and IT fluxes across plasma membrane were investigated. The concentration of enzymes for protoplast isolation, e.g. cellulase, pectolyase, macerozyme, hemicellulase, and sorbitol content, incubation time were systemically studied. High yield and viability of protoplast was achieved after 6-8 hours incubation of P. euphratica callus in enzyme solution containing 1.5% （w：v） cellulase R-10, 0.1% （w：v） pectolyase Y-23, 0.2% （w：v） macerozyme R-10, 0.05% （w：v） hemicellulase and 0.75M）.80 mol·L^-1 sorbitol. Non-invasively ion selective microelectrode technique was used to access proton fluxes in the absence and presence of NaCl （20 mmol.L-1）. Salt-induced transient net IT effiux was observed in the plasma membrane ofP. euphratica cells. The shift of IT flux response to NaC1 shock and the relevance to salt tolerance were discussed.

Retrieval of leaf biochemical properties by inversed PROSPECT model and hyperspectral indices:an application to Populus euphratica polymorphic leaves

Leaf biochemical properties have been widely assessed using hyperspectral reflectance information by inversion of PROSPECT model or by using hyperspectral indices, but few studies have focused on arid ecosystems. As a dominant species of riparian ecosystems in arid lands, Populus euphratica Oliv. is an unusual tree species with polymorphic leaves along the vertical profile of canopy corresponding to different growth stages. In this study, we evaluated both the inversed PROSPECT model and hyperspectral indices for estimating biochemical properties of P. euphratica leaves. Both the shapes and biochemical properties of P. euphratica leaves were found to change with the heights from ground surface. The results indicated that the model inversion calibrated for each leaf shape performed much better than the model calibrated for all leaf shapes, and also better than hyperspectral indices. Similar results were obtained for estimations of equivalent water thickness （EWT） and leaf mass per area （LMA）. Hyperspectral indices identified in this study for estimating these leaf properties had root mean square error （RMSE） and R2 values between those obtained with the two calibration strategies using the inversed PROSPECT model. Hence, the inversed PROSPECT model can be applied to estimate leaf biochemical properties in arid ecosystems, but the calibration to the model requires special attention.

Differences in chlorophyll fluorescence parameters and water content in heteromorphic leaves of Populus euphratica from Inner Mongolia, China

We studied three typical heteromorphic leaves of Populus euphratica trees growing in the Wuhai region of Inner Mongolia,China,i.e.,lanceolate,broad-ovate and dentate broad-ovate leaves and mainly focused on the changes in chlorophyll fluorescence parameters and free water and bound water content.The results show that the values of Fm（maximal fluorescence yield）,Fv/Fm（maximum photochemical quantum yield of PSII） and Fv/F0（potential quantum efficiency of PSII） of lanceolate leaves were the least on young trees,while these parameters were the least on the ovate leaves of old trees.Compared with young trees,the free water content of heteromorphic leaves of old trees increased significantly,i.e.,by 78.94% in lanceolate leaves and in the leaves of broad-ovate and dentate broad-ovate by 10.99% and 10.60%,respectively.Correlation analysis showed that free water content is significantly related to Fv/Fm and Fv/F0 in young trees,while the relationship of total water content with Fv/Fm and Fv/F0 is positive in old trees.

Ultrastructural and Extracellular Protein Changes in Cell Suspension Cultures of Populus euphratica Associated with Low Temperature-induced Cold Acclimation

Populus euphratica Olive is the only tree species that can grow in the saline land and also survive cold winters in northwest China, and it plays a very important role in stabilizing the vulnerable ecosystem there. A cell suspension culture was initiated from callus derived from plantlets of Populus euphratica. Cold acclimation was induced (LT50 of 17.5 ℃) in cell suspension at 45 ℃ in the dark for 30 days and the freezing tolerance increased from LT50 of 12.5 ℃ in nonacclimated cells to LT50 of 17.5 ℃ in cold-acclimated cells. Microvacuolation, cytoplasmic augmentation and accumulation of starch granules were observed in cells that were cold-acclimated by exposure to low temperatures. Several qualitative and quantitative changes in proteins were noted during cold acclimation. Antibodies to carrot extracellular (apoplastic) 36 kD antifreeze protein did not cross react on immunoelectroblots with extracellular proteins in cell suspension culture medium of Populus euphratica, indicating no common epitopes in the carrot 36 kD antifreeze protein and P. euphratica extracellular proteins. The relationship of these changes to cold acclimation in Populus euphratica cell cultures was discussed.

Plant diversity and its maintenance in Populus euphratica riparian forests in the Ejina Oasis,China

The dynamic environments in riparian zones support a variety of life-history strategies, which constitute a fundamental mechanism for development and maintenance of biodiversity in riparian forest ecosystems. To demonstrate the effect of life-history strategies on biodiversity, we investigated community-level diversity and its relationship to environmental variability in the riparian Populus euphratica forests of the Ejina Oasis. Communities were divided into 14 associations on the basis of their species impor- tance values. The Simpson＇s index, Shannon-Wiener index, Pielou＇s evenness index and the regional Whittaker＇s index were applied to calculate community diversity. An ordination of the sample plots was carried out by correspondence analysis （CA）. Biodiversity was relatively low across the entire study area, but there was high community diversity （flw = 8.09） due to the spatial heterogeneity of habitats in different plots. In addition, we investigated the relationship between biodiversity and several environmental factors, such as water availability, community components and soil conditions. We conclude that the heterogeneity of soil and water availability drives community diversity patterns in riparian zones and that community-level diversity favors the maintenance of species diversity in the P euphratica forests in the Ejina Oasis.

Sap-flow measurement and scale transferring from sample trees to entire forest stand of Populus euphratica in desert riparian forest in extreme arid region

Understanding how the transpiration of this vegetation type responds to environmental stress is important for determining the wa-ter-balance dynamics of the riparian ecosystem threatened by groundwater depletion. Transpiration and sap flow were measured using the heat-pulse technique. The results were then projected up to the stand level to investigate the stand’s water-use in relation to climate forcing in the desert riparian forest in an extreme arid region. This study took place from April through October 2003 and from May through October 2004. The experimental site was selected in the Populus euphratica Forest Reserve (101o10' E, 41o59' N) in Ejina county, in the lower Heihe River basin, China. The sapwood area was used as a scalar to extrapolate the stand-water consumption from the whole trees’ water consumption measured by the heat-pulse velocity recorder (HPVR). Scale transferring from a series of individual trees to a stand was done according to the existing natural variations between trees under given environmental conditions. The application of the biometric parameters available from individual tree and stand levels was proved suitable for this purpose. A significant correlation between the sapwood area and tree diameter at breast height (DBH) was found. The prediction model is well fitted by the power model. On the basis of the prediction model, the sapwood area can be cal-culated by DBH. The sap-flow density can then be used to extrapolate the stand-water use by means of a series of mathematical models.

Measuring and modeling evapotranspiration of a Populus euphratica forest in northwestern China

Vegetation is sparsely distributed in the arid regions of northwestern China,and accurately measuring and partitioning of evapotranspiration is of importance for ecosystems in such areas.In this study,we measured and analyzed diurnal and phenological variations in evapotranspiration using the eddy-covariance method based on the Penman-Monteith,Priestley-Taylor,Shuttleworth-Wallace models,a newly proposed improved dual source model and a clumped model in a forest reserve in the Ejin oasis of Populus euphratica in 2015 and 2016 growing seasons.A sensitivity analysis was performed for the models with higher accuracies and we examined the biotic and abiotic controls on evapotranspiration.The results show that the total amounts of evapotranspiration during the two growing seasons in 2015 and 2016 were 622 and 612 mm,respectively.Phenological variations in evapotranspiration produced single-peak curves,while diurnal variations reflected the influence of high temperatures on some afternoons.The Priestley-Taylor and the improved dual source models gave the most accurate evapotranspiration values at the daily scale and appeared to be most suitable for the estimation of evapotranspiration for the species in arid regions.In addition,both models were the most sensitive to net radiation(Rn).

Short-term transcriptomic responses of Populus euphratica roots and leaves to drought stress

Populus euphratica Oliv.,the Euphrates poplar,is the tallest tree species in the arid desert areas of Northwest China.Investigation of its drought-resistance genes is valuable to increase understanding of drought resistance mechanisms.RNA-seq of leaves and roots under drought simulation by 25%polyethylene glycol-6000(PEG 6000)were performed at 0,4,and 12 h.Leaves and roots responded differently to drought via differentially upregulated and downregulated genes;more genes were downregulated than upregulated in both leaves and roots.Additionally,these differentially expressed genes were enriched in different GO terms and KEGG pathways.For example,GO term‘response to organic substance’and KEGG pathway‘nitrogen metabolism’were enriched for drought-stressed leaves,while GO term‘cell wall organization or biogenesis’and KEGG pathway‘zeatin biosynthesis’were enriched for drought-stressed roots.The enrichment of the GO term‘enzyme linked receptor protein signalling pathway’in both leaf and root drought responses suggests that these tissues may also have similar mechanisms.However,roots under drought stress for four hs responded by activating programed cell death.The KEGG pathway‘plant hormone signal transduction’was detected for 4-and 12-h droughtstressed leaves and 12-h drought-stressed roots,suggesting that plant hormone signal transduction plays an important role in both roots and leaves.GO enrichment of upregulated and downregulated genes for leaves and roots reflect differentially regulatory mechanisms of response to drought stress via different biological processes such as the regulation of photosynthesis and auxin signalling pathway in leaves,and the regulation of defence response and water homeostasis in roots.Fifteen candidate genes,including transcription factors,protein kinase,transporter,late embryogenesis abundant protein and mannitol dehydrogenase,were further selected to determine their response to drought using qRTPCR.The results show that the expression patterns of 13 of the 15 genes correspond to the RNA-seq results.This study provides new insights into the drought response mechanism of P.euphratica and suggests new candidate gene resources.

Spatial pattern and utility of Populus euphratica seed rain:a simulation study combined with field investigations

Seed dispersal is a fundamental process affecting destinies of seeds and seedlings, as well as regeneration dynamics and distribution patterns of communities. Recently however, declining Populus euphratica forests have failed to regenerate by seeding, while the spatial pattern and utility of their seed rains are unclear. In this study, we designed a model to simulate seed rains based on field investigations by fitting our observations to a normal distribution and testing the model with data from three years, with ac- ceptable results. We examined the simulated results of dispersed seeds patterns on the classification of three factors, i.e., density of Sophora alopecuroides, surface soil moisture and surface soil salinity. The results of seed rain simulation show that over 70% of seeds were dispersed and confined to each of the three plots where their mother trees located. The proportion of 3：7 seeds dispersed inside and outside each plots remained largely unaltered. The differences in the amounts of dispersed seed among the different levels of each of the factors were not significant, although the distributing pattern of P. euphratica in each plot varied. Therefore, in P. eu- phratica communities, the amount of seed does not become a constraint in subsequent germination, although the surface environment does. We conclude that successful P. euphratica seed regeneration relies on less than 30% of seeds dispersed over longer distance to colonize favorable growth habitats.

Construction of a cDNA-AFLP reaction system of Populus euphratica for abiotic stress studies

We established a cDNA-AFLP （amplified fragment length polymorphism） system for Populus euphratica, after optimizing several key factors. The results show that optimized CTAB method is suitable for the extraction of ideal RNA from P euphratica; the double-stranded cDNA （ds-cDNA） is completely digested by the combination of EcoRI and MseI for 4.5 h; the reducible result can be obtained when the preamplification products were diluted 10-fold for selective amplification templates. The amplification products resulting from this reaction system were determined by silver staining and the fragments were clear and stable. Detected by RT-PCR, the optimized differential display cDNA-AFLP technique can accurately reflect the true differences of gene expressions among plants. In addition, 42 pairs of primer arrays, producing repeatable and prolific bands, were successfully selected by this improved method. This investigation establishes a cDNA-AFLP system for P. euphratica, which forms a basis for further study on stress-response genes in P. euphratica.

Cloning of plasma membrane H^+-ATPase gene in Populus euphratica Oliv.

For this paper, the plasma membrane （PM） H^＋-ATPase gene has been cloned from Populus euphratica Oliv. through a ho- mology based strategy. The isolated 3,210 bp cDNA contains a single 2,862 bp open reading frame （ORF） which encodes a putative H^＋-ATPase protein of 953 amino acid residues, with a significant homology to plasma membrane H^＋-ATPase of Prunus persica, Phaseolus vulgaris, Sesbania rostrata and Daucus carota. The predicted protein has a molecular weight of 104,553 Da. The copy number analysis revealed multiple copies of the PM H^＋-ATPase in the P. euphratica genome after digestion of their genomic DNA by the restriction enzymes EcoRI, NdeI, FbaI and Bg/Ⅱ, and Southern blot.

Influence of environmental,root,and stand parameters on soil surface CO_2 efflux in a Populus euphratica of desert forest in extreme arid region

Spatial variation in soil surface CO2 efflux was measured in a stand of Populus euphratica in the Ejina Oasis of desert riparian forest in the extreme arid region in northwestern China from April 2007 through October 2007.Measurements were taken with a gas-exchange analyzer linked to a soil-respiration chamber.The mean soil CO2 efflux in the stand was 2.71 μmol/(m2·s) during the growing season and 1.38 μmol/(m2·s) in the nongrowing season.The seasonal maximum (end of May through early June) andminimum (October) CO2 efflux were 3.38 and 0.69 μmol/(m2·s),respectively.The diurnal fluctuation of CO2 efflux was relatively small (< 20 percent),with theminimum appearing around 05:00 and the maximum around 15:00.Linear regression analysis showed soil-surface CO2 efflux to be most highly correlated with soil temperature (R2=0.435) and soil moisture (R2=0.213).When all variables were considered simultaneously,only soil temperature (R2=0.378),soil moisture (R2=0.147),and root volume density (R2=0.021) explained a significant amount of variance in soil surface CO2 efflux.Stand volumes were not correlated with soil CO2 efflux on our sites.