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Research areaThe YREB covers Shanghai, Jiangsu, Zhejiang, Anhui, Jiangxi, Hubei, Hunan, Chongqing, Sichuan, Guizhou, and Yunnan. It includes the Yangtze River Delta urban agglomerations (YRDUA), Yangtze River midstream urban agglomeration (YRMUA), and Chengdu-Chongqing urban agglomeration (CCUA). With a regional area of 2.05 million km2, the YREB runs through the eastern, central and western regions in China32. In 2019, the total GDP of YREB is 45.8 trillion yuan, accounting for 46.2% of the national GDP. The YREB plays a pivotal strategic support and leading role in the overall situation of stable economic growth in China33. At the same time, the contradiction between the shortage of land resources and economic growth in the YREB is very prominent. Therefore, this paper selects 107 cities in YREB as the research sample. The specific geographic locations are shown in Fig. 2. This article uses ARCGIS 10.2 version to draw the map. The URL link is http://demo.domain.com:6080/arcgis/services.Figure 2The geographic location of the YREB in China.Full size imageResearch methodsGlobal Malmquist–Luenberger indexUGLUE refers to the effective utilization degree of land elements under certain input of other elements. The green utilization of urban land mainly comes from three aspects: first, improve the utilization intensity of the existing actual input land, that is, increase the input intensity of other elements of the unit land area. Second, reduce the input of land in the production process to avoid excessive waste of land. Third, promote the optimal allocation of land elements among production units. Technical efficiency refers to the maximum degree that all factor inputs need to expand or shrink in equal proportion when all production units reach the production frontier. However, for production units with high technical efficiency, the factor allocation structure may not be reasonable. The land factors may still have the problem of under-input or over-input, resulting in the reduction of UGLUE.Pastor and Lovell34 proposed a global index, which uses all the inspection periods of each decision-making unit as a benchmark to construct the production frontier. According to the current benchmark construction period t, the production possibility set reference set is defined as follows:$$P_{C}^{t} (x^{t} ) = left{ {left. {(y^{t} ,b^{t} )} right|x^{t} {kern 1pt} can{kern 1pt} , produce{kern 1pt} , b^{t} ,y^{t} } right}$$
(1)
The global benchmark is defined as: (P_{G} = P_{C}^{1} , cup ,P_{C}^{2} , cup , cdots ,P_{C}^{t}), The subscripts C and G represent the current benchmark and the global benchmark respectively. The ML index of decision-making unit i is calculated based on the current reference benchmark:$$ML^{S} (x^{t} ,y^{t} ,b^{t} ,x^{t + 1} ,y^{t + 1} ,b^{t + 1} ) = frac{{1 + D_{C}^{S} (x^{t} ,y^{t} ,b^{t} )}}{{1 + D_{C}^{S} (x^{t + 1} ,y^{t + 1} ,b^{t + 1} )}}$$
(2)
Among them, the superscript S indicates two adjacent periods, t period and t + 1 period. The subscript C indicates the current benchmark, which is a simplified directional distance function. (ML^{s} > 1), indicates that the productivity increases. (ML^{s} < 1), indicates that the productivity decreases.According to Hofmann et al.35, the GMLI is defined as follows:$$GMLI^{t,t + 1} (x^{t} ,y^{t} ,b^{t} ,x^{t + 1} ,y^{t + 1} ,b^{t + 1} ) = frac{{1 + D_{G}^{T} (x^{t} ,y^{t} ,b^{t} )}}{{1 + D_{G}^{T} (x^{t + 1} ,y^{t + 1} ,b^{t + 1} )}}$$ (3) Among them, (D_{G}^{T} (x,y,b) = max left{ {alpha |(y - alpha y,b - alpha b) in P_{G} (x)} right}). (GMLI^{t,t + 1} > 1) indicates that the productivity has increased. (GMLI^{t,t + 1} < 1) indicates that the productivity decreases. The GMLI is further broken down as follows:$$begin{aligned} & GMLI^{t,t + 1} (x^{t} ,y^{t} ,b^{t} ,x^{t + 1} ,y^{t + 1} ,b^{t + 1} ) = frac{{1 + D_{G}^{T} (x^{t} ,y^{t} ,b^{t} )}}{{1 + D_{G}^{T} (x^{t + 1} ,y^{t + 1} ,b^{t + 1} )}} \ & quad = frac{{1 + D_{G}^{t} (x^{t} ,y^{t} ,b^{t} )}}{{1 + D_{G}^{t + 1} (x^{t + 1} ,y^{t + 1} ,b^{t + 1} )}} times left[ {frac{{(1 + D_{G}^{T} (x^{t} ,y^{t} ,b^{t} ))/(1 + D_{C}^{T} (x^{t} ,y^{t} ,b^{t} ))}}{{(1 + D_{G}^{T} (x^{t + 1} ,y^{t + 1} ,b^{t + 1} ))/(1 + D_{C}^{T} (x^{t + 1} ,y^{t + 1} ,b^{t + 1} ))}}} right] \ & quad = frac{{TE^{t + 1} }}{{TE^{t} }} times left( {frac{{BPG_{t + 1}^{t + 1} }}{{BPG_{t}^{t + 1} }}} right) = EC_{t}^{t + 1} times BPC_{t}^{t + 1} \ end{aligned}$$ (4) Among them, TE is the change of technological progress. EC is the change of technological efficiency. The change of technological progress reflects the change of the highest technical level. The improvement of the highest technical level often requires the introduction and innovation of advanced technology, which often requires a large amount of investment. The change of technical efficiency reflects the gap with the highest technical level. Narrowing the gap with the highest technical level often requires improvements in internal management and governance structures. (BPG_{t}^{t + 1}) is the “best practitioner gap” between the current period and overall technological frontier. (BPC_{t}^{t + 1}) measures the changes in the “best practitioner gap” between two periods (technological changes). (BPC_{t}^{t + 1} , > , 1 ,) indicates technological progress. (BPC_{t}^{t + 1} < 1) indicates technology regress.Econometric techniques of industrial agglomeration on UGLUEIn recent years, many scholars used the traditional SPM for empirical analysis, which is a basic measurement model suitable for panel data. Therefore, this article firstly uses the traditional SPM to analyze the impact of industrial agglomeration on UGLUE. The formula is:$$begin{aligned} ln UGLUE_{it} & = alpha_{0} + alpha_{1} ln RZI_{it} + alpha_{2} ln RZI_{it} *ln RZI_{it} + alpha_{3} ln RDI_{it} + alpha_{4} ln EC_{it} \ & quad + alpha_{5} ln GDP_{it} + alpha_{6} ln TEC_{it} + alpha_{7} ln ROAD_{it} + alpha_{8} ln GOV_{it} + varepsilon_{it} \ end{aligned}$$ (5) Among them, ε is the disturbance term. i represents the city, i in this paper involves 107 cities in YREB. t represents the time, and the range of t in this paper is from 2007 to 2016. UGLUE is the explained variable, which represents the UGLUE. RZI and RDI are explanatory variables, representing industrial specialization agglomeration and industrial diversification agglomeration. EC is the industrial structure. GDP is the level of economic development. TEC is the level of technology. ROAD is the level of infrastructure. GOV is the degree of government intervention. (alpha_{1}) to (alpha_{8}) is the coefficient of each variable.Formula (5) assumes that the UGLUE changes with the changes of various influencing factors in the current period. That is, there is no time lag effect. But in reality, land use often has a time lag effect. The previous level has a non-negligible impact on the current results. Therefore, this paper selects the dynamic panel model for empirical analysis. However, there is often a two-way causal relationship between industrial agglomeration and UGLUE, which may cause endogenous bias. For example, cities with higher UGLUE levels tend to have higher levels of economic development, which promotes industrial agglomeration in this city. Therefore, this paper adopts the method of system GMM for regression analysis of dynamic panel model. Compared with mixed OLS, system GMM can make full use of sample information, select appropriate lag terms as instrumental variables36. It can effectively solve the endogeneity problem between industrial agglomeration and UGLUE. Based on the above analysis, this paper introduces the first-order lag term of UGLUE on the basis of formula (5). The DPM is as follows:$$begin{aligned} ln UGLUE_{it} & = beta_{0} + tau ln UGLUE_{i(t - 1)} + beta_{1} ln RZI_{it} + beta_{2} ln RZI_{it} times ln RZI_{it} + beta_{3} ln RDI_{it} \ & quad + beta_{4} ln EC_{it} + beta_{5} ln GDP_{it} + beta_{6} ln TEC_{it} + beta_{7} ln ROAD_{it} + beta_{8} ln GOV_{it} + varepsilon_{{{text{it}}}} \ end{aligned}$$ (6) Among them, (tau) is the first-order lag coefficient of UGLUE, reflecting the time lag effect of UGLUE.Variable descriptionExplained variableThe GMLI is used to measure the UGLUE of 107 cities in YREB. According to existing research37, the following core evaluation index of UGLUE are selected (see Table 1). Regarding input indicators, we mainly choose land element input M, labor element input L, and capital element input K as input indicators. Regarding output indicators, we choose the added value of the secondary and tertiary industries in the municipal area as the expected output, and use the GDP deflator to convert it into a comparable value. At the same time, pollution indicators such as industrial wastewater emissions, industrial sulfur dioxide emissions, and industrial smoke (dust) emissions are selected as undesired output. Since the GMLI reflects the growth rate of UGLUE, this paper assumes that the GMLI in 2006 is 1, and then multiplies the calculated GMLI year by year to obtain the development level of UGLUE in each city from 2007 to 2016.Table 1 Input and output index.Full size tableExplanatory variablesIndustrial specialization index ZI is usually used to measure the specialization level of urban industries. The specialization index is represented by the share of the employment of the industry in the total employment of the city:$$ZI_{i} = Max_{j} (S_{ij} )$$ (7) Nextly, we use the relative specialization index to make a horizontal comparison of the specialization level between different cities:$$RZI_{i} = Max(S_{ij} /S_{j} )$$ (8) The most common measure of the level of industrial diversification is the Herfindahl–Hirschman Index (HHI). For city i, the HHI is the sum of the square sum of employment shares of all industries in the city. The diversification index is the reciprocal of the HHI:$$DZ_{i} = frac{1}{{sumlimits_{j} {S_{ij}^{2} } }}$$ (9) The expression of relative diversification index is as follows:$$RDI_{i} = {1 mathord{left/ {vphantom {1 {sumlimits_{j} {left| {S_{ij} - S_{j} } right|} }}} right. kern-0pt} {sumlimits_{j} {left| {S_{ij} - S_{j} } right|} }}$$ (10) Among them, Sij is the employment proportion of j industry in city i, and Sj is the proportion of the total employment of the national j industry. The greater value of RZI and RDI, the higher level of industrial specialization and diversification.Control variablesRegarding control variables, we choose the following variables as control variables.Industrial structure (EC): The continuous optimization of industrial structure promotes the improvement of UGLUE through three aspects: saving land, increasing land income and promoting the optimal allocation of land resources. This paper selects the added value of the tertiary industry as a percentage of GDP (take the logarithm) to express.Technological level (TEC): The higher the technological innovation level of a city is, the more it promotes the use of input elements and the transformation of innovation results, thereby improving the UGLUE. This paper selects the proportion of science and technology expenditure to fiscal expenditure (take the logarithm) to represent.Economic development level (GDP): The continuous economic development promote the rational allocation of various production factors and increase the level of urban land output, thereby improving the UGLUE. This paper selects GDP per capita (take the logarithm) to express.Road infrastructure level (ROAD): The continuous improvement of infrastructure reduces transportation costs and transaction costs, and promotes communication externalities between producers, consumers, and between producers and consumers. This paper selects the average road area per capita (take the logarithm) to express.Government behavior (GOV): Fiscal expenditure is an important means for the government to carry out macro-control. Appropriate fiscal expenditure makes up for market shortages, improves factor flow and resource allocation efficiency, and realizes positive economic externalities. This paper selects the proportion of fiscal expenditure to GDP (take the logarithm) to express. We can see the meaning of specific variables from Table 2.Table 2 The descriptive statistics of variables.Full size tableData sourceThe object of this thesis is the 107 cities in YREB from 2007 to 2016. The urban construction land area data comes from the "China Urban Construction Statistical Yearbook", and the rest of the index data all come from the "China City Statistical Yearbook". The URL link is https://www.cnki.net/. In order to maintain the integrity of the data, this article uses the average method to fill in the missing values. In addition, because Chaohu City began to be under the jurisdiction of Hefei City in 2011, Bijie City and Tongren City in Guizhou Province only became prefecture-level cities in 2011. The three cities and Pu'er City are taken from the sample to maintain the continuity of data. More

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Study siteThe field experiment was conducted at Namco Station (30°47’N, 90°58’E, altitude 4730 m) of the Institute of Tibetan Plateau Research, Chinese Academy of Sciences (ITPCAS), which is located in the alpine steppes of TP in China. The experiment was permitted by ITPCAS, complied with local and national guidelines and regulations. From 2006 to 2017, the mean annual temperature (MAT) and mean annual precipitation (MAP) was about − 0.6 °C and 406 mm, respectively. Monthly mean temperature varied from − 10.8 °C in January to 9.1 °C in July and most of the precipitation occurred from May to October37,38. During our six-year observations (2010, 2011, 2012, 2013, 2015 and 2017), climate change during the growing season from May to September varied differently, with the annual precipitation ranged from 255.9 mm to 493.8 mm and the MAT from 6.7 to 7.4 °C. Androsace tapete, Kobresia pygmaea, Stipa purpurea and Leontopodium pusillum were the dominant plant species at the alpine steppe.Experimental design and treatmentsThe long-term experiment began in May, 2010. Three homogenous plots were randomly arranged as replicates at the alpine steppe and six subplots (~ 13 m2) were distributed in each plot by a cycle, with a 2 m buffer zone between each adjacent subplot (Appendix S1: Fig. S1). In this experiment, six treatments of N fertilization rate (0, 1, 2, 4, 8, and 16 g N m−2 yr−1) were clockwise applied in each subplot. The subplots of 0 g N m−2 yr−1 were control group. We sprayed NH4NO3 solution on the first day of each month in the growing season (from May to September) each year. After fertilizing, we rinsed plant residual fertilizer with a little deionized water (no more than 2 mm rainfall). For the control groups, we added equivalent amount of water. The experiment was conducted from 2010 to 2017 (it should be pointed out that there was no fertilization in 2014 and 2016).Sampling and measurementsThe samples were collected with the training and permission of ITPCAS and involved plants that are common species and not endangered or protected. The identification of the plants was done by referring to a book of Chen and Yang39. Pictures of the corresponding specimens can be seen on the website of ITPCAS (http://itpcas.cas.cn/kxcb/kxtp/nmc_normal_plant/).Vegetation samples were collected in August in 2011 and repeated at the same time in 2012, 2013, 2015 and 2017. We established one 50 × 50 cm quadrat in each subplot, clipped aboveground biomass (AGB) and sorted species by families. The biomass was used to measure ANPP (g m−2 yr−1). Following aboveground portion collected, we used three soil cores (5 cm diameter) to collect the belowground roots at 0–30 cm depth and mixed into one sample, which were used to assess belowground net primary productivity (BNPP, g m−2 yr−1). The roots were cleaned with running water to remove sand and stones.Both plant and root samples were dried at 75 °C for 48 h and then ground into powder (particle size ~ 5 μm) by a laboratory mixer mill (MM400, Retsch). To determine N and C content of plants, we weighed the samples into tin capsules and measured with the elemental analyzer (MAT253, Finnigan MAT GmbH, Germany).Estimation of the critical N rate (Ncr), N retention fraction (NRF), N retention capacity and N-induced C gainAccording to the N saturation hypothesis, plant productivity increases gradually during N addition, reaches a maximum at the Ncr, and eventually declines16,17. We considered the Ncr to be the rate where ANPP no longer remarkably changed with N addition (Fig. 1).We defined plant N retention fraction (NRF, %; Eq. 1) as the aboveground N storage caused by unit N addition rate, and N retention capacity (g N m−2 yr−1; Eq. 2) was the increment of N storage due to exogenous N addition compared to the control40. The equations are as following:$$N;retention;fraction = frac{{ANPP_{tr} times N;content_{tr} – ANPP_{ck} times N;content_{ck} }}{N;rate}$$
(1)
$$N;retention;capacity = ANPP_{tr} times N;content_{tr} – ANPP_{ck} times N;content_{ck}$$
(2)
where ANPPtr and N contenttr (%) refer to those in the treatment (tr) groups, and ANPPck and N contentck refer to those in the control (ck) groups. These expressions are also used in the following equations (Eqs. 3–5).The N-induced C gain (g C m−2 yr−1; Eq. 3) was estimated by the increment of C storage owing to exogenous N addition compared to the control40. Maximum N retention capacity (MNRC, Eq. 4) and maximum N-induced C gain (Eq. 5) mean the maximum N and C storage increment in plant caused by exogenous N input at Ncr, respectively. The formulas are as following:$$N{text{-}}induced;C;gain = ANPP_{tr} times C;content_{tr} – ANPP_{ck} times C;content_{ck}$$
(3)
$$MNRC = ANPP_{max } times N;content_{max } – ANPP_{ck} times N;content_{ck}$$
(4)
$$Maximum;N{text{-}}induced;C;gain = ANPP_{max } times C;content_{max } – ANPP_{ck} times C;content_{ck}$$
(5)
where ANPPmax, N contentmax and C contentmax refer to the value of ANPP, N content and C content at Ncr, respectively.Data synthesisTo evaluate N limitation and saturation on the TP more accurately, we searched papers from the Web of Science (https://www.webofscience.com) and the China National Knowledge Infrastructure (https://www.cnki.net). The keywords used by article searching were: (a) N addition, N deposition or N fertilization, (b) grassland, steppe or meadow. Article selection was based on the following conditions. First, the experimental site must be conducted in a grassland ecosystem. Second, the experiment had at least three N addition levels and a control group. Third, if the experiment lasted for many years, we analyzed data with multi-year average. Based on the above, we collected 89 independent experimental cases. Among these, 27 cases were located on the TP alpine grasslands, 25 in the Inner Mongolia (IM) grasslands and 37 in other terrestrial grasslands (detailed information sees Appendix S2: Table S1).We extracted ANPP data and N addition rate of these cases and estimated Ncr and ANPPmax (Appendix S2: Fig. S2). We then calculated NRF, N retention and C gain of each group of data for further analysis (Appendix S2: Table S2). Most of the 89 cases did not have data on N and C content. To facilitate the calculation, we summarized N and C content from 40 articles in the neighboring areas of the cases and divided the N and C content into seven intervals according to the N addition rate (Appendix S2: Table S3 and Fig. S3). The unit of N addition rate was unified to “g N m−2 yr−1”. All the original data were obtained directly from texts and tables of published papers. If the data were displayed only in graphs, Getdata 2.20 was used to digitize the numerical data. For the estimation of N retention and C gain of the TP at current N deposition rates and future at Ncr, we fitted the exponential relationship to the data from 27 cases on the TP, and then substituted N rates into the fitted equations (Eq. 6):$$y = a times left[ {1 – exp left( { – bx} right)} right].$$
(6)
We also included MAT, MAP, soil C:N ratio, fencing management (fencing or grazing) and grassland type (meadow, steppe and desert steppe) of the experiment sites for exploring the drivers affecting N limitation (Appendix S2: Table S1). When climatic data were missing from the article, MAT and MAP were obtained from the WorldClim (http://www.worldclim.org).Species were usually divided into four functional groups (grasses, sedges, legumes and forbs) to study the response of species composition to N addition in previous study41. We synthesized 13 TP experimental cases (including our field experiment) from the data synthesis and each case included at least three functional groups (detailed references see Appendix S2).Statistical analysisThere were 42 species in our field experiment. We divided them by family into eleven groups: Asteraceae (forbs), Poaceae (grasses), Leguminosae (legumes), Rosaceae (forbs), Boraginaceae (forbs), Caryophyllaceae (forbs), Cyperaceae (sedges), Labiatae (forbs), Primulaceae (forbs), Scrophulariaceae (forbs) and Others. Due to species in the group of Others contributed only 1.22% of AGB, we analyzed AGB and foliar stoichiometry among other ten families (Appendix S1: Table S1). In Namco steppe, forbs, grasses, sedges and legumes accounted for 78.0%, 7.4%, 8.2% and 5.2% of the AGB respectively (Appendix S1: Table S1 and Fig. S2). Such a large number of forbs suggested that our experiment was conducted on a severely degraded grassland.For our field data, two-way ANOVAs were used to analyze the effects of year, N fertilization rate and their interactions on species AGB. One-way ANOVAs were used to test the response of ANPP, BNPP, root:shoot ratio, species foliar C content, N content and C:N ratio to N addition rate. Duncan’s new multiple range test was used to compare the fertilization influences at each rate in these ANOVAs. Prior to the above ANOVAs, we performed homogeneity of variance test and transformed the data logarithmically when necessary. Simple regression was used to estimate the relevance among ANPP, NRF, N retention capacity and C gain with N addition rates.Structural equation modeling (SEM) was used to explore complex relationships among multiple variables. To quantify the contribution of drivers such as climate and soil to Ncr, ANPP, NRF and MNRC, we constructed SEM based on existing ecological knowledge and the possible relationships between variables. We considered environmental factors (MAT, MAP and soil C:N) and ANPPck as explanatory variables, and Ncr, NRF and MNRC as response variables. We included the ANPPck in the SEM rather than the ANPPmax because we wonder whether there was a relationship between ANPP in the absence of exogenous N input and the ecosystem N retention in the presence of N saturation. This has important implications for assessing N input. Before constructing the SEM, we excluded collinearity between the factors. In addition, Student’s t-test and one-way ANOVAs were performed to explain the effect of fencing management and grassland type on above response variables, respectively. The SEM was constructed using the R package “piecewiseSEM”42. Fisher’s C was used to assess the goodness-of-model fit, and AIC was for model comparison.Given the influence of extreme values in the data synthesis, we calculated the geometric mean of Ncr, NRF, N retention and N-induced C gain. All statistical analyses were performed with SPSS 26.0 and RStudio (Version 1.2.1335) based on R version 3.6.2 (R Core Team, 2019). More