Patterns and driving factors of leaf C,N,and P stoichiometry in two forest types with different stand ages in a mid-subtropical zone

Background:Carbon(C),nitrogen(N),and phosphorus(P)stoichiometry is a key indicator of nutrient utilization in plants,and C/N/P ratios are related to the life histories and adaptation strategies of tree species.However,no consensus has been reached on how leaf stoichiometric characteristics are affected by forest type and stand ages.The relationships between leaf stoichiometry and geographical,meteorological,and soil factors also remain poorly understood.Methods:Leaf and soil were sampled from forest stands of different age groups(young,middle-aged,near-mature,and mature)in two forest types(Chinese fir(Cunninghamia lanceolata)forests and evergreen broadleaved forests).The relationships between leaf C,N,and P stoichiometric parameters and geographical,meteorological,and soil factors were analysed by using redundancy analysis(RDA)and stepwise linear regression analysis.Results:Leaf C concentrations peaked in the near-mature stands with increasing age irrespective of forest type.Leaf N and P concentrations fluctuated with a rising trend in Chinese fir forests,while decreased first and increased later from young to mature phases in natural evergreen broadleaved forests.Chinese fir forests were primarily limited by N and P,while natural evergreen broadleaved forests were more susceptible to P limitation.Leaf C,N,and P stoichiometric characteristics in Chinese fir forests were mainly affected by the soil total P concentration(SP),longitude(LNG),growing season precipitation(GSP)and mean temperature in July(JUT).The leaf C concentration was mainly affected by GSP and JUT;leaf N and P concentrations were both positively correlated with LNG;and leaf P was positively correlated with SP.In evergreen broadleaved forests,however,leaf stoichiometric parameters displayed significant correlations with latitude(LAT)and mean annual precipitation(MAP).Conclusions:Leaf stoichiometry differed among forest stands of different age groups and forest types.Leaf C,N,and P stoichiometry was primarily explained by the combinations of SP,LNG,GSP and JUT in Chinese fir forests.LAT and MAP were the main controlling factors affecting the variations in the leaf C,N,and P status in natural evergreen broadleaved forests,which supports the temperature-plant physiological hypothesis.These findings improve the understanding of the distribution patterns and driving mechanisms of leaf stoichiometry linked with stand age and forest type.

Afforestation with an age-sequence of Mongolian pine plantation promotes soil microbial residue accumulation in the Horqin Sandy Land, China

Land use change affects the balance of organic carbon(C)reserves and the global C cycle.Microbial residues are essential constituents of stable soil organic C(SOC).However,it remains unclear how microbial residue changes over time following afforestation.In this study,16-,23-,52-,and 62-year-old Mongolian pine stands and 16-year-old cropland were studied in the Horqin Sandy Land,China.We analyzed changes in SOC,amino sugar content,and microbial parameters to assess how microbial communities influence soil C transformation and preservation.The results showed that SOC storage increased with stand age in the early stage of afforestation but remained unchanged at about 1.27-1.29 kg/m2 after 52 a.Moreover,there were consistent increases in amino sugars and microbial residues with increasing stand age.As stand age increased from 16 to 62 a,soil pH decreased from 6.84 to 5.71,and the concentration of total amino sugars increased from 178.53 to 509.99 mg/kg.A significant negative correlation between soil pH and the concentration of specific and total amino sugars was observed,indicating that the effects of soil acidification promote amino sugar stabilization during afforestation.In contrast to the Mongolian pine plantation of the same age,the cropland accumulated more SOC and microbial residues because of fertilizer application.Across Mongolian pine plantation with different ages,there was no significant change in calculated contribution of bacterial or fungal residues to SOC,suggesting that fungi were consistently the dominant contributors to SOC with increasing time.Our results indicate that afforestation in the Horqin Sandy Land promotes efficient microbial growth and residue accumulation in SOC stocks and has a consistent positive impact on SOC persistence.

Changes in leaf stomatal traits of different aged temperate forest stands

Stomata control carbon and water vapor exchange between the leaves and the atmosphere,thus infl uencing photosynthesis and transpiration.Combinations of forest patches with different stand ages are common in nature,however,information of which stomatal traits vary among these stands and how,remains limited.Here,seven different aged forest stands(6,14,25,36,45,55,and 100 years)were selected in typical temperate,mixed broadleaf-conifer forests of northeast China.Stomatal density,size and relative area of 624 species,including the same species in stands of different ages were selected.Stomatal density,size and relative area were distributed log-normally,differing across all species and plant functional groups.Stomatal density ranged from 4.2 to 1276.7 stomata mm^(–2),stomatal size ranged from 66.6 to 8315.7μm^(2),and stomatal relative area 0.1–93.3%.There was a significant negative relationship between density and size at the species and functional group levels,while the relative stomatal area was positively correlated with density and size.Stomatal traits of dominant species were relatively stable across different stand ages but were significantly different for herbs.The results suggest that stomatal traits remain relatively stable for dominant species in natural forests and therefore,spatial variation in stomatal traits across forest patches does not need to be incorporated in future ecological models.

Stem sap flow of Haloxylon ammodendron at different ages and its response to physical factors in the Minqin oasis-desert transition zone, China

Haloxylon ammodendron, with its tolerance of drought, high temperature, and salt alkali conditions, is one of the main sand-fixing plant species in the oasis-desert transition zone in China. This study used the TDP30(where TDP is the thermal dissipation probe) to measure hourly and daily variations in the stem sap flow velocity of H. ammodendron at three age-classes(10, 15, and 20 years old,which were denoted as H10, H15, and H20, respectively) in the Minqin oasis-desert transition zone,China, from May through October 2020. By simultaneously monitoring temperature, relative humidity,photosynthetically active radiation, wind speed, net radiation, rainfall, and soil moisture in this region, we comprehensively investigated the stem sap flow velocity of different-aged H. ammodendron plants(H10,H15, and H20) and revealed its response to physical factors. The results showed that, on sunny days, the hourly variation curves of the stem sap flow velocity of H. ammodendron plants at the three age-classes were mainly unimodal. In addition, the stem sap flow velocity of H. ammodendron plants decreased significantly from September to October, which also delayed its peak time of hourly variation. On rainy days, the stem sap flow velocity of H. ammodendron plants was multimodal and significantly lower than that on sunny days.Average daily water consumption of H. ammodendron plants at H10, H15, and H20 was 1.98, 2.82, and 1.91kg/d, respectively. Temperature was the key factor affecting the stem sap flow velocity of H. ammodendron at all age-classes. Net radiation was the critical factor influencing the stem sap flow velocity of H.ammodendron at H10 and H15;however, for that at H20, it was vapor pressure deficit. The stem sap flow velocity of H. ammodendron was highly significantly correlated with soil moisture at the soil depths of 50and 100 cm, and the correlation was strengthened with increasing stand age. Altogether, our results revealed the dynamic changes of the stem sap flow velocity in different-aged H. ammodendron forest stands and its response mechanism to local physical factors, which provided a theoretical basis for the construction of new protective forests as well as the restoration and protection of existing ones in this region and other similar arid regions in the world.

Comparison of transpiration between different aged black locust(Robinia pseudoacacia)trees on the semi-arid Loess Plateau,China

Black locust （Robinia pseudoacacia） is widely planted throughout the semi-arid Loess Plateau of China. The spatial distribution of this species at different ages is highly heterogeneous due to restoration and management practices. In this study, we aimed to compare the transpiration levels between different aged black locusts at the tree and stand scales, clarifying the physiological status of this species with different ages. Black locust trees with two representative age classes （12 and 28 years） were selected in the Yangjuangou catchment on the semi-arid Loess Plateau. Sap flux density （Fd） and environmental variables （solar radiation, air temperature, relative humidity and soil water content） were simultaneously monitored throughout the growing season of 2014. Tree transpiration （Et） was the product of Fd and sapwood area （As）, and stand transpiration （Ec） was calculated basing on the stand sap flux density （Js） and stand total sapwood area （AsT）. Stomatal conductance （gs） was measured in a controlled environment and hydraulic conductance was estimated using the relationship between transpiration rate and vapor pressure deficit （VPD）. Our results showed that Et and Ec were higher in the 28-year-old stand than in the 12-year-old stand. The gs and hydraulic conductance of 28-year-old trees were also higher than those of 12-year-old trees, and the two parameters were thus the causes of variations in transpiration between different age classes. After rainfall, mean Fd increased by 9% in 28-year-old trees and by 5% in 12-year-old trees. This study thus suggests that stand age should be considered for estimating transpiration at the catchment and region scales in this area. These results provide ecophysiological evidences that the older black locust trees had more active physiological status than the younger ones in this area. These findings also provide basic information for the management of water resources and forests on the semi-arid Loess Plateau.

Biomass and Nutrient Accumulation Characteristics of Young Stands of Alnus cremastogyne

To investigate plant biomass and nutrient distribution and accumulation in organs of Alnus cremastogyne at different ages from 1 to 4 years, the biomass, N, P, K, Ca, Mg, Fe and Zn were tested. The results showed that the average biomass of the whole tree and the biomass of leaf, branch, stem, and root were in positive correlation with tree age, but the growth rate of biomass had a decreasing trend with the tree age increasing, and only the biomass proportion of the trunk in the whole individual plant showed an increasing trend with age. The contents of nutrient elements in organs showed an order of N 〉 Ca 〉 K 〉 Mg 〉 P 〉 Fe 〉 Zn ; and the contents of N, P and K were higher in the leaf than in other organs, and the contents of Ca, Mg and Fe in the root were higher than in other organs. The accumulations of N, P, K, Ca, Mg and Zn were the highest in the trunk, and that of Fe was the highest in the root. The annual net accumulations of N, P, K, Ca and Mg in the average trees from 1 to 4 years old were 17.07, 40.79, 95.82 and 106.71 g, respectively, and the annual net accumulations of microelements （Fe and Zn） were 335.04, 577.26, 1267 and 1525.27 mg, respectively.

Carbon sequestration in biomass and soil following reforestation:a case study of the Yangtze River Basin

The effect of reforestation on carbon sequestration has been extensively studied but there is less understanding of the changes that stand age and vegetation types have on changes in biomass carbon and soil organic carbon(SOC)after reforestation.In this study,150 reforested plots were sampled across six provinces and one municipality in the Yangtze River Basin(YRB)during 2017 and 2018 to estimate carbon storage in biomass and soil.The results illustrate that site-averaged SOC was greater than site-averaged biomass carbon.There was more carbon sequestered in the biomass than in the soil.Biomass carbon accumulated rapidly in the initial 20 years after planting.In contrast,SOC sequestration increased rapidly after 20 years.In addition,evergreen species had higher carbon density in both biomass and soil than deciduous species and economic species(fruit trees).Carbon sequestration in evergreen and deciduous species is greater than in economic species.Our findings provide new evidence on the divergent responses of biomass and soil to carbon sequestration after reforestation with respect to stand ages and vegetation types.This study provides relevant information for ecosystem management as well as for carbon sequestration and global climate change policies.

Soil microbial attributes along a chronosequence of Scots pine(Pinus sylvestris var. mongolica) plantations in northern China

Soil microorganisms play a key role in soil organic matter dynamics, nutrient cycling, and soil fertility maintenance in forest ecosystems, and they are influenced by stand age and soil depth. However, few studies have simultaneously considered these two factors. In this study, we measured soil microbial biomass carbon (SMBC), soil microbial biomass nitrogen (SMBN), soil basal respiration (SBR) rate, and potential extracellular enzyme activity (EEA) in soil to a depth of 60 cm under 10-, 30-, and 40-year-old Scots pine (Pinus sylvestris var. mongolica) stands (Y10, Y30, and Y40, respectively) in plantations in northern China in 2011. Soil water content (SWC), soil pH, soil organic carbon (SOC), and soil total nitrogen (STN) were also measured to explore their effects on soil microbial indices across different stand ages and soil depths. Our results showed that SMBC, SMBN, and the SBR rate were generally higher for the Y30 stand than for the Y10 and Y40 stands. Potential EEA, except forα-glucosidase, decreased significantly with increasing stand age. Soil organic carbon,STN, SWC, and soil pH explained 67%of the variation in soil microbial attributes among the three stand ages. For the same stand age, soil microbial biomass and the SBR rate decreased with soil depth. Lower microbial biomass, lower SBR rate, and lower EEA for the mature Y40 stand indicate lower substrate availability for soil microorganisms, lower soil quality, and lower microbial adaptability to the environment. Our results suggest that changes in soil quality with stand age should be considered when determining the optimum rotation length of plantations and the best management practices for afforestation programs.

Contrasting effects of N addition on the N and P status of understory vegetation in plantations of sapling and mature Larix principis-rupprechtii

Aims The productivity of forest plantations in temperate areas is often lim-ited by nitrogen(N),but may shift towards phosphorus(P)limitation with increasing atmospheric N deposition.Nutrient resorption is a nutrient conservation strategy in plants.Although data on nutrient resorption are available for overstory trees,there are few data for understory vegetation.Methods We examined leaf N and P concentrations and N and P resorption efficiencies(NRE and PRE,respectively)in eight understory species in 11-and 45-year-old Larix principis-rupprechtii stands subjected to N supplementation over a 3-year period.Important findings Leaf N concentrations and N:P ratios increased and P concentrations decreased,with N input in species within the 45-year-old stand,but not in the 11-year-old stand.NRE and PRE were not altered by N input in any of the species in either stand,but N resorption pro-ficiency decreased and P resorption proficiency increased,in the species in the 45-year-old stand.Thus,the growth of understory species may be more P-limited in the 45-versus 11-year-old stand,and nutrient resorption proficiency was more sensitive to N add-ition than nutrient resorption efficiency.These results will improve the understanding of nutrient use strategies and their responses to N addition in understory vegetation.The contrasting effects of N addition on nutrient status between stand ages cannot be ignored when modeling ecosystem nutrient cycling under global N depos-ition conditions.