Studies on the Secretion of Organic Acids from Roots of Two-year-old Larix olgensis under Nutrient and Water Stress

[Objective] This study aimed to investigate the effects of nutrient and water stress on the secretion of organic acids from roots of two-year-old Larix olgensis. [Method] Different nutrient and water stress conditions were designed to investi- gate the types and quantities of organic acids secreted from roots of two-year-old L. olgensis. [Result] Under nutrient and water stress, the types of organic acids secreted from roots of two-year-old L. olgensis increased, which varied with different stress levels. In addition, nutrient and water stress also increased the secretory vol- ume of organic acids from roots of two-year-old L. olgensis. The increment in total amount of organic acids reached the maximum under Level 1 （high stress）. Among different types of organic acids, the increment in secretory volume of citric acid reached the maximum, followed by malic acid, while oxalic acid presented relatively small increment. Furthermore, the increment in secretory volume of these three organic acids all reached the maximum under Level 1. [Conclusion] The proportion of the secretory volume of each organic acid accounting for the total amount of organic acids varied slightly, but the overall order was unchanged.

Family selection and evaluation of Larix gmelinii var.principis-rupprechtii(Mayr.)Pilger based on stem analysis data at multiple sites

Larix gmelinii var.principis-rupprechtii(Mayr.)Pilger is an important native tree species in North China with advantages of fast growth,straight trunk,and good wood properties.The multi-year and multi-site breeding research of families of the species has not been reported previously.Based on diameter at breast height(DBH),height and volume of 25 families on four experimental sites,we calculated variance components,genetic parameters,juvenile and mature trait correlations and made genotype main effect plus genotype×environment interaction effect(GGE)biplot based on the breeding values estimated using the method of best linear unbiased prediction(BLUP).Compared with height,DBH and volume had higher heritability and larger variation coefficients,making them the more suitable traits for family selection and evaluation.Based on these,GGE biplots containing 20 combinations of site×age were drawn using data at 13 to 17 years when the interactions between family and location were strong.Test sites classifications based on DBH,and volume were inconsistent,with two categories for DBH and one for volume.The Guyuan site was the most suitable with strong discriminating ability,high representativeness and stability among tree ages.Integrating the ranking results of DBH and volume,families 66,76,82 and 111 were high-yielding and stable,families 78 and96 were high-yielding with above average stability,families72 and 79 were high-yielding with below average stability,whereas stability of family 100 was inconsistent between DBH and volume.Early selection based on DBH was convenient and reliable,and can be made at seven years.This study provides support for the selection of Larix gmelinii var.principis-rupprechtii families in Hebei province and an example for the application of stem analysis data from multiple sites in tree breeding.

Nutrient retranslocation from the fine roots of Fraxinus mandshurica and Larix olgensis in northeastern China

Nutrient retranslocation in trees is important in nutrient budgets and energy flows in forest ecosystems. We investigated nutrient retranslocation in the fine roots of a Manchurian Ash （Fraxinus mandshurica） and a Larch （Larix olgensis） plantation in northeastern China. Nutrient retranslocation in the fine roots was investigated using three methods, specifically, nutrient concentration, the ratio of Ca to other elements （Ca/other elements ratio） and nutrient content. The method based on nutrient content proved most suitable when investigating nutrient retranslocation from fine roots of the two species. The nutrient-content-based method showed that there were retranslocations of N, P, K and Mg from the fine roots of Manchurian Ash, with retranslocation efficiencies of 13, 25, 65, and 38 %, respectively, whereas there were no Ca retranslocations. There were retranslocations of N, P, K, Ca and Mg from the fine roots of Larch, with retranslocation efficiencies of 31, 40, 52, 23 and 25 %, respectively.

Changing regularity of plant and insect community characteristics due to pest outbreaks in Larix principis- rupprechtii plantations

Research has indicated that simple forest eco-system composition,structure and diversity have uncompli-cated community relationships and insufficient pest control capabilities.To investigate changing characteristics of plant and insect communities in under pest outbreaks in Larix principis-rupprechtii plantations,the research areas were defined as mature(48–50 years)and young(24–29 years)infested stands along with healthy stands.The results show a reduction in the complexity and diversity of plant communi-ties and herbaceous plant guilds(polycultures of beneficial plants)and the complexity and dominance of insect com-munities,especially natural insect enemies.The results also show the relative simplicity of the main factors of commu-nity change and development that represent the characteris-tics of pest outbreaks in L.principis-rupprechtii plantations.The complexity and diversity of plant communities,particu-larly herbaceous plant guilds play a fundamental role in the regulation and development in forest ecosystems.

Phenological observations on Larix principis-rupprechtii Mayr. in primary seed orchard

Through 5 years of phenological observations on Larix principis-rupprechtii Mayr. in primary seed orchard and studies on population and individuals of clones, the annual periodic phenological laws were revealed and the annual phe-nological periodic table was drawn up. The correlation between various phenophases, the air temperature and active accumu-lated temperature were analyzed and expounded. The authors also analyzed the similarities and differences of phenophases among clonal individuals as well as the blooming properties of male and female flowers at the same time. This study could pro-vide theoretical reference for working out the production plan of improved varieties and other management measures in seed orchard of Larix principis-rupprechtii.

Spatial distribution of fine roots of larch and ash in the mixed plantation stand

The spatial distribution of standing fine roots in tree rows of different species in a 12-year-old mixed stand of ash (Fraxinus mandshurica Rupr.) and larch (Larix olgensis Henry) was studied by soil core sampling in early spring, 2001. It is found that ash and larch differ greatly in their belowground biomass distribution. Ash has much higher fine root biomass density in the soil than larch at stand level (with the max value of 4442.3 vs. 2234.9 gm-3). Both tree species deployed more fine roots in their neighboring zone, suggesting a less intensive competition between roots of the two species. Both fine root biomass density and root length density of ash in the zone between larch tree rows are greater than that of larch in zone between ash tree rows, indicating that ash is more powerful than larch in belowground competition. The spatial distribution feature of roots favors the growth of ash in the mixed stand.

Age–age correlations and early selection for growth traits in 40 half-sib families of Larix principis-rupprechtii

Larix principis-rupprechtii Mayr is a dominant species in coniferous forests of North China.However,early selection of L.principis-rupprechtii for growth traits is poorly characterised.To explore the optimal selection age for this species,heights(HT)and diameters at breast(DBH)of 40 half-sib families were measured at ages 3,12,22,and 28 years in a progeny test population established in the town of Kangjiahui,Shanxi Province.Age trends in heritability,age–age genetic correlations,and early selection efficiency for height and DBH were analysed.The individual heritability of these growth traits varied over time,and maximized at different ages(0.55 at age 12 for HT and 0.48 at age 28 for DBH).The age–age genetic correlations were always positive,and the majority were high(0.790–0.953)between the juvenile and mature ages for HT and DBH.For the same pairs of measurements,HT demonstrated higher age–age genetic correlations than DBH,and both age–age genetic correlation data sets were described well by the linear relationship with the logarithm of the age ratio(r2[0.90).The regression slope for DBH was lower than that for HT.Based on the early selection efficiency estimates,the optimal selection age could be as early as age 6 for DBH and 8–9 years for HT.The results of this study provide information that can be used to assist early selection practices in L.principis-rupprechtii improvement programs in Shanxi Province.

Variations in stem radii of Larix principis-rupprechtii to environmental factors at two slope locations in the Liupan Mountains,northwest China

Relationships between stem growth and climatic and edaphic factors,notably air temperatures and soil moisture for different slopes,are not completely understood.Stem radial variations were monitored at the bottom and top slope positions in a Larix principis-rupprechtii plantation during the 2017 and 2018 growing seasons.Total precipitation during the growing season in 2017 and 2018 was 566 mm and 728 mm,respectively.Stem contractions typically occurred after mid-morning followed by swelling in the late afternoon in both plots,reflecting the diurnal cycle of water uptake and loss.Trees at the two locations showed the same growth initiation(mid-May)because of the small differences in air and soil temperatures.There were no significant differences in cumulative stem radial growth between the bottom plot(1.57±0.34 mm)and the top plot(1.55±0.26 mm)in 2018.However,in 2017,the main growth period of the bottom plot ceased 17 days earlier than in the top plot,while cumulative seasonal growth of the bottom plot(1.08±0.25 mm)was significantly less than the top plot(1.54±0.43 mm).Maximum daily stem shrinkage was positively correlated with air and soil temperatures,solar radiation,vapor pressure deficits,and negatively correlated with volumetric soil moisture content.The maximum daily shrinkage reflected transpiration rates as affected by environmental factors.Daily radial stem increment was correlated with precipitation and volumetric soil moisture in both years,but with air temperatures only in 2017.The seasonal growth of L.principis-rupprechtii Mayr thus shows interannual dynamics,while precipitation constitutes a key driving factor.

Estimation of leaf area index from high resolution ZY-3 satellite imagery in a catchment dominated by Larix principis-rupprechtii,northern China

Leaf area index(LAI) is a key factor that determines a forest ecosystem's net primary production and energy exchange between the atmosphere and land surfaces.LAI can be measured in many ways, but there has been little research to compare LAI estimated by different methods. In this study, we compared the LAI results from two different approaches, i.e., the dimidiate pixel model(DPM) and an empirical statistic model(ESM) using ZY-3 high-accuracy satellite images validated by field data. We explored the relationship of LAI of Larix principis-rupprechtii Mayr plantations with topographic conditions. The results show that DPM improves the simulation of LAI(r = 0.86,RMSE = 0.57) compared with ESM(r = 0.62, RMSE =0.79). We further concluded that elevation and slope significantly affect the distribution of LAI. The maximum peak of LAI appeared at an aspect of east and southeast at an elevation of 1700–2000 m. Our results suggest that ZY-3 can satisfy the needs of quantitative monitoring of leaf area indices in small-scale catchment areas. DPM provides a simple and accurate method to obtain forest vegetation parameters in the case of non-ground measurement points.

Rules of Changes in Soil Nutrients and Enzyme Activities of Larix principis-rupprechtii in Different Forest Ages

In this study,three different ages of Larix principis-rupprechtii forests in 5,10 and 20 years were selected as the research objects,and the changes in soil nutrient and soil enzyme activities in different growth stages were analyzed. The results showed that the contents of organic matter and available phosphorus in the soil of different growth stages showed a significant downward trend with the increase of soil depth.For different forest ages in the same soil layer,the soil available phosphorus content declined with the increase of the forest age. The organic matter content of 40-60 cm in 20 years of forest age was the lowest in July,which was 4. 17 g/kg,significantly lower than that in other soil layers. Besides,the soil available phosphorus content of 5 years of forest age reached the maximum in July,with an average of 4. 44 mg/kg,which was higher than the available phosphorus content in soil in May and September,but the difference between the three months was not significant. The changes in ammonium nitrogen and nitrate nitrogen content in soil with different forest ages were consistent with the changes in the soil depth,showing a downward trend. In the new leaf stage,the ammonium nitrogen content of the L. principis-rupprechtii forest land in5 years of age at 20-40 and 40-60 cm of the soil depth was 13. 47 and 9. 09 mg/kg,respectively,which was 46. 9% and 64. 2% lower than that at 0-20 cm( 25. 36 mg/kg) of the soil depth. The soil nitrate nitrogen content of 20 years of forest age was 19. 24 mg/kg,which was25. 8% lower than that of 10 years of forest age,showing significant difference( P < 0. 05). In addition,with the increase of the age of L. principis-rupprechtii,soil catalase( CAT),alkaline phosphatase( ALP) and urease( Ure) decreased,and the decline of ALP was slow,while CAT and Ure decreased significantly. In summary,it is concluded that the soil fertility of forest land declined with the increase of forest age on the basis of change trend of soil nutrient and soil enzyme activity in the surveyed forest age.

Effect of thinning intensity on the stem CO_(2) efflux of Larix principis-rupprechtii Mayr

Background:Stem CO_(2) efflux(E_(S))plays a critical role in the carbon budget of forest ecosystems.Thinning is a core practice for sustainable management of plantations.It is therefore necessary and urgent to study the effect and mechanism of thinning intensity(TI)on E_(S).Methods:In this study,five TIs were applied in Larix principis-rupprechtii Mayr 21-,25-,and 41-year-old stands in North China in 2010.Portable infrared gas analyzer(Li-8100 A)was used to measure ES and its association with environmental factors at monthly intervals from May to October in 2013 to 2015.In addition,nutrients,wood structure and nonstructural carbon(NSC)data were measured in August 2016.Results:The results show that ES increased with increasing TI.The maximum ES values occurred at a TI of 35%(3.29,4.57 and 2.98μmol·m^(-2)·s^(-1))and were 1.54-,1.94-and 2.89-fold greater than the minimum E_(S) value in the CK stands(2.14,2.35 and 1.03μmol·m^(-2)·s^(-1))in July for the 21-,25-and 41-year-old forests,respectively.The E_(S) of the trees in low-density stands was more sensitive to temperature than that of the trees in high-density stands.Soluble sugars(SS)and temperature are the main factors affecting ES.When the stand density is low enough as 41-year-old L.principis-rupprechtii forests with TI 35%,bark thickness(BT)and humidity should be considered in addition to air temperature(T_(a)),wood temperature(T_(w)),sapwood width(SW),nitrogen concentration(N)and SS in the evaluation of ES.If a change in stand density is ignored,the CO_(2) released from individual 21-,25-and 41-year-old trees could be underestimated by 168.89%,101.94% and 200.49%,respectively.CO_(2) release was estimated based on the stem equation in combination with the factors influencing ES for reference.Conclusions:We suggest that it is not sufficient to conventional models which quantify ES only by temperature and that incorporating the associated drivers(e.g.density,SS,SW and N)based on stand density into conventional models can improve the accuracy of ES estimates.

Range-wide genetic diversity in natural populations of Larix principis-rupprechtii Mayr.

Prince Rupprecht’s larch(Larix principis-rupprechtii Mayr.),a deciduous conifer,widely grows in middle and high elevations of Northern China.Its natural distribution has sharply decreased and has become fragmented,which may have resulted in the loss of genetic variation.In this study,ten natural populations across the entire range of this species were analyzed using amplifi ed fragment length polymorphism markers.A total of 309 loci were detected from 225 individuals of these populations,of which 261(84.5%)were polymorphic.At the species level,the genetic diversity was high(average of the Nei’s genetic diversity H e=0.2602,and Shannon’s information index I=0.3967).The results of molecular variance analysis showed that 90.71%of the genetic diversity occurred within populations.The genetic diff erentiation among populations was moderate as a whole(F ST=0.0929,G ST=0.1510),which is consistent with the moderate level of gene fl ow among populations(N m=2.8116).Based on the unweighted pair group method with arithmetic mean and STRU CTU RE analysis,these populations were grouped into three genetically distinct clusters.The degree of inter-population diff erentiation(G ST=0.1338)for the south group was larger than that for the north group(G ST=0.0915).There was a signifi cant correlation between genetic distance and geographic distance across the species range(r=0.316,P<0.05).Genetic diversity was signifi-cantly associated with longitude but not elevation or climatic factors.The populations with high genetic diversity from each cluster are therefore recommended for future conservation and management of this species.

Characterization of expressed sequence tags obtained by SSH during root formation in Larix cuttings

SSH was used to analyze gene transactivation during root formation of Larix cuttings. Two subtractive cDNA libraries were constructed from clone 31-6 as tester or driver and clone 15-4 as driver or tester. The SSH PCR products from the libraries were cloned into a pGEM-T easy vector and after PCR and dot blot analysis, positive clones were selected, sequenced and compared to the database in GenBank with BLASTX. The results of a sequence assembly in two libraries show that 521 UniEST （expressed sequence tag） were obtained. These 521 UniEST belong to metabolism, signal pathways, transport, resistance, developmental processes, local- ization, unknown proteins and ＂no hits found＂. All of these suggest that subtractive cDNA libraries during root formation of Larix cuttings were constructed successfully.

扫描根窗法在落叶松人工林细根动态研究中的应用

植物根系深藏于地下,由于缺乏有效的观测方法,制约了对根系的了解。以落叶松人工林为研究对象,根据扫描根窗法对落叶松人工林细根长期观测案例,介绍扫描根窗法在野外原位观测细根动态的过程。结果表明:在深秋季节安装根窗,根窗安装扰动后恢复期为1 a,经过冻融交替和一个生长季后,细根现存量迅速恢复且趋于稳定;通过观察发现吸收根现存量峰值多出现在春季和初夏,而运输根峰值多出现在秋季;野外根窗法与微根管法相比可清晰的观测到细根的完整分枝结构,将高度异质性的细根群体细分为不同根序或功能类型,为区分不同细根功能和分枝等级提供了手段。

More tree growth reduction due to consecutive drought and its legacy effect for a semiarid larch plantation in Northwest China

Extreme climate has increasingly led to negative impacts on forest ecosystems globally,especially in semiarid areas where forest ecosystems are more vulnerable.However,it is poorly understood how tree growth is affected by different drought events.In 2006–2009,the larch plantations in the semiarid areas of Northwest China were negatively affected by four consecutive dry years,which was a very rare phenomenon that may occur frequently under future climate warming.In this study,we analyzed the effect of these consecutive dry years on tree growth based on the data of the tree rings in the dominant layer of the forest canopy on a larch plantation.We found that the tree-ring width index(RWI)in dry years was lower than that in normal years,and it experienced a rapidly decreasing trend from 2006 to 2009(slope=-0.139 year^(-1),r=-0.94)due to water supply deficits in those dry years.Drought induced legacy effects of tree growth reduction,and consecutive dry years corresponded with greater growth reductions and legacy effects.Growth reductions and legacy effects were significantly stronger in the third and fourth consecutive dry years than that of single dry year(p<0.05),which might have been due to the cumulative stress caused by consecutive dry years.Our results showed that larch trees experienced greater tree growth reduction due to consecutive dry years and their legacy effect,and the trees had lower recovery rates after consecutive dry years.Our results highlight that consecutive dry years pose a new threat to plantations under climate warming,and thus,the effect of climate extremes on tree growth should be considered in growth models in semiarid areas.

Spacial Patterns of Fine Root Abundance in Mixed Larch-Ash Plantation

Horizontal and veftical distributions of fine root abundance (mass per unit ground surface arca) were investi-gated in a densely planted larch - ash -stripe - mixed tbrest on dark broxvn forest soil in northeast China. There was evidencefor some degree of below-ground niche partitioning (or differentiation) betmpen the two species in hoth the horizontal andvertical directions. The ash fine roots largely penetrated into the larch belt (larch Sub-community) in surttrce soil (0-20 cm),indicating a possible inductive effect of larch ecological field on ash roots, while the penetration of larch tine roots into ashbelt (ash sub-community) was much restricted- which reflected a negative influence of ash ccological tield on Iarch roots. Inthe vertical direction of marginal soil the ash tine roots were mainly distributed in topsoil with a vertical gradient similar tothat as in the internal ash sub-community. but the larch fine roots were relatively compelled to deepcr soil layers by the com-petition (or exclusion) of marginal ash trees. All the dit1brences or complemcntarity wcre considered to the result of inter-specitic competition, which was important to the coexistence of the two tbrest species and the sustainability of mixed planta-tion.

Discrepancy in fine root turnover estimates between diameter-based and branch-order-based approaches:a case study in two temperate tree species

Fine root turnover plays a key role in carbon （C） budgets and nutrients cycles in forest ecosystems. However, the difference between branch-order-based and diameter-based approaches in estimating fine root turnover is still unclear. We studied root biomass turnover based on multiplying root standing biomass by turnover rate （inverse of median root longevity） in two Chinese temperate tree species, Fraxinus mand- shurica Rupr. and Larix gmelinii Rupr. The minirhizotron （MR） tech- nique was used to estimate longevities for first and second order roots, and total roots （R,o,al） apparent on the MR tube surface. The correspond- ing biomass for each root group was estimated by soil monolith. The difference in biomass turnover between Rtotal and the sum of the first and second order roots was used to represent the discrepancy between di- ameter- and order-based approaches. First order roots had shorter life spans and higher biomass turnover rates than the second order roots in both species. Biomass turnover estimated by the order-based method for F. mandshurica and L. gmelinii were 155.4 g.m2.aland 158.9 g.m2.a-1, respectively, in comparison with 99.5 g.m^-2-a^-1 and 117.7 g.m^-2.a^-1 estimated by the diameter-based method, indicating that the diameter-based approach underestimated biomass turnover. The most probable reason was that the order-based method enhanced separation of the heterogeneous root population into relatively homogenous root groups with varying turnover rates. We conclude that separating fine root pool into different branch orders can improve the accuracy of estimates for fine root turnover, as well as the understanding of the belowground C allocation and nutrient cycling at ecosystem level.

不同火烧强度对兴安落叶松根系分解和土壤有机碳的影响

细根分解作为陆地化学循环的关键环节,也是土壤养分的重要来源。为了解火干扰对细根分解的影响,以内蒙古大兴安岭毕拉河自然保护区兴安落叶松林为研究对象,利用分解袋法分析火烧后土壤理化性质、根系质量残留率和营养元素变化情况,以及驱动根系分解的关键因子。结果表明:①火烧后细根分解速率显著加快。轻度与重度火烧后,极细根(d<0.5 mm)细根质量残留率分别减少了10.15%和8.14%;中等细根(0.5 mm≤d<2.0 mm)在重度火烧处理下,细根质量残留率减少了4.78%;粗根(d≥2.0 mm)在重度火烧处理下,根系质量残留率显著增加了5.29%。②短期内,火烧降低了土壤有机碳质量分数,土壤化学计量特性是影响细根分解的主要因子,而细根碳质量分数始终是影响火烧前后细根分解的关键因子,其对分解的贡献率均在20%以上,轻度火烧条件下极细根碳在分解中的贡献率达到23.72%。③火烧对兴安落叶松(Larix gmelinii)根系分解和土壤有机碳具有积极影响,且根系分解对土壤有机碳库具有显著作用,其总效应达到0.3。因此,火烧通过影响土壤化学性质加快了兴安落叶松细根分解速率,并且轻度火烧对根系分解产生的影响较大。

不同林龄华北落叶松人工林叶凋落物分解及养分动态比较

采用网袋法,对0～540 d内不同林龄华北落叶松(Larix principis-rupprechtii)人工林叶凋落物的分解特征及营养元素质量分数变化动态进行了比较分析。结果表明:经过540 d的分解,幼、中、近、成熟林4个龄组的叶质量损失率为48.47%～61.72%,并呈极显著差异,不同林龄的叶凋落物分解速率呈现出成熟林>近熟林>中龄林>幼龄林。用含哑变量的Olson指数衰减模型对叶凋落物的分解动态进行拟合,结果显示,幼龄林、中龄林、近熟林和成熟林叶凋落物的分解系数为0.028 30～0.036 19;半衰期和周转期分别为1.59～2.04 a和6.90～8.82 a。在整个分解过程中,4个龄组的养分质量分数变化一致,随着时间的延长,N质量分数不断升高;P质量分数呈现出先降低后上升的趋势;K、C质量分数和C/N呈下降的趋势。在分解过程中,4个龄组的N、P、K、C的释放率在总体上变化一致,C和K不断释放,且为净释放状态,呈现直接释放模式;N的释放率为富集-释放模式;P的释放率表现为淋溶-富集-释放模式。结果表明,成熟林和近熟林的叶凋落物比幼龄林和中龄林的叶凋落物易分解,且成熟林和近熟林中的营养元素也较易释放。

施肥对日本落叶松人工林细根直径、根长和比根长的影响

以辽宁东部山区16年生日本落叶松人工林为研究对象,探讨施肥对日本落叶松1～5级根序中细根直径、根长和比根长的影响.结果表明：随着根序等级的增加,日本落叶松细根平均直径和根长显著增加（P＜0.05,P＜0.01）、比根长则显著下降（P＜0.01）.在日本落叶松的5级根序中,1级根的平均直径最细、根长最短、比根长最高,而5级根则相反;随着根序等级的增加,日本落叶松细根平均直径、根长和比根长的变异系数逐渐增大.除1级根外,土层对细根的平均直径、根长和比根长没有显著影响（P＞0.05）.与对照样地相比,施肥对各级细根平均直径、根长和比根长的影响主要表现在1～2级根上,对3级根序以上的细根影响不显著（P＞0.05）.其中,施氮肥显著降低了1～2级根的平均直径（P＜0.05）,施氮肥以及氮磷肥显著降低了表层土壤（0～10 cm）中1级根的平均根长（P＜0.05）,表层土壤中细根的比根长在施氮肥的条件下显著增加（P＜0.05）.