Evaluation of the growth, adaption, and ecosystem services of two potentially-introduced urban tree species in Guangzhou under drought stress
Study site, tree selections, and drought-simulation experimentThis research was performed in Guangzhou (22°26′-23°56′N, 112°57′-114°03′E), which is a core city located in subtropical zones. With an area of 7434.4 km2 and a population of 18.87 million, Guangzhou’s urbanization rate has reached 86.46%. To cope with multiple environmental challenges, several urban-forest nurseries were established to cultivate and introduce various tree species. Among them, we selected the one in Tianhe District as our study site. This nursery was not only abundant with native and exotic tree species but also equipped with similar edatope in cities, which was ideal for our research.Tilia cordata Mill. (Tc) and Tilia tomentosa Moench (Tt), originating from the west of Britain and southeast of Europe, were common urban tree species planted in European cities. Based on their performance in providing ecological and landscape functions, these two tree species were considered to be introduced for urban greening. Therefore, Tilia cordata Mill. (Tc) and Tilia tomentosa Moench (Tt) were selected as our objectives, which were investigated for their growth and ecosystem services to evaluate their adaption in Guangzhou. In addition, a native tree species Tilia miqueliana Maxim (Tm) was also implemented concurrent measurement as a comparison.For each of the three surveyed tree species, ten trees with a diameter at breast height (DBH) around 5.5 cm and tree height around 2.5 m were chosen for our experiment, which were thought to possess similar initial statuses. To investigate the impact of drought on the growth and ecosystem services of the three selected tree species, a controlled experiment was launched from January to December in 2020. For each tree species, five trees were planted in the common environment as the controlled group, while the other five trees were under the precipitation-exclusion installation (PEI) as the drought-simulation group. Consisting of several water-proof tents, PEI was adequately large and could completely prevent trees from obtaining rainfalls, which created a precipitation-exclusive environment to simulate an enduring drought event within the whole research period (Fig. 1).Figure 1Schematic diagram of the drought simulation experiment for the three surveyed tree species.Full size imageEnvironmental monitoring systemsClimatic data were sampled every 10 min with a weather station (WP3103 mesoscale automatic weather station, China) located at an unshaded site in the nursery. The data were stored in the logger and copied to our laboratory to produce daily or monthly data. All the climatic variables, including photosynthetically active radiation (PAR, µmol m-2 s-1), wind speed (m s-1), precipitation (mm), and air temperature (°C) were calculated from January to December in 2020.For volumetric soil water content (%; VWC), the HOBO MX2307 system (Onetemp, Adelaide, Australia), placed in a shaded box in the nursery, was applied for all the three tree species from both the controlled and drought-simulation groups. For each individual tree, the sensing probe was inserted horizontally at the depths of 30 cm and located 20 cm in the northern direction from the tree stems. Based on the daily readings, monthly means were calculated from January to December in 2020.Measurement of above-ground growthTo investigate the above-ground growth of the three tree species from both the controlled and drought-simulation groups, their DBH (diameter at breast height, cm), tree height (m), and LAI (leaf area index) were measured at the beginning of each month in 2020. DBH was measured with the help of a caliper (Altraco Inc., Sausalito, California, USA), and their tree heights were measured using a standard tape. The crown analytical instrument CI-110 (Camas, Washington State, USA) was used to capture an accurate image of tree crowns and calculate LAI. Sufficient numbers of points were measured and recorded to describe each tree’s average crown shape. The software FV2200 (LICOR Biosciences, Lincoln, NE) helped compute each tree’s crown width and crown area.Measurement of below-ground growthFine root coring campaigns were launched for all the trees of the three tree species from both the controlled and drought treatment groups every three to four months, i.e., in February, May, September, and December. Although the coring campaign might damage part of the roots, the fine roots obtained each time were a mere portion of the whole root system, not affecting the general development of trees’ underground processes. For every individual tree, two 30-cm soil cores were applied in each direction of north, south, east, and west, of which one was located at 20 cm to the trunk (paracentral roots) and the other one was located at 40 cm (outer roots). In addition, the soil samples were evenly divided into three horizons which were 0–10 cm (shallow layer), 10–20 cm (middle layer), and 20–30 cm (deep layer). Then a sieve with 2-mm mesh size was used to filter all the fine roots. The fine roots were washed carefully to remove the adherent soils and dried in an oven at 65 ℃ for 72 h. Finally, all the samples were weighed using a balance with an accuracy of four decimal places to obtain the dry weight. The fine root biomass at different depths was calculated using the dry weight divided by the cross-sectional area of the auger20.Model’s simulation of ecosystem servicesThe process-based model City-Tree was used to predict the ecosystem services of the three tree species from both the controlled and drought-simulation groups23. The model required the data of tree growth parameters including tree height, DBH, and crown area together with environmental conditions such as edaphic and climatic data24. In this research, cooling, evapotranspiration and CO2 fixation of the three surveyed tree species in the controlled and drought-treatment groups were simulated at the end of 2020.The actual evapotranspiration eta was calculated from the potential evapotranspiration using fetp[t], Tilia’s factors fetp[t], and the reduction factor fred:$${mathrm{et}}_{mathrm{a}}={mathrm{f}}_{mathrm{red}}*{mathrm{f}}_{mathrm{etp}}left[mathrm{t}right]*{mathrm{et}}_{mathrm{p}}$$The process of tree’s evapotranspiration (etp) was calculated on the basis of SVAT algorithm together with Penman formula in the module on water balance as below:$${mathrm{et}}_{mathrm{p}}=left[mathrm{s }/ left(mathrm{s}+upgamma right)right]*left({mathrm{r}}_{mathrm{s}}-{mathrm{r}}_{mathrm{L}}right) /mathrm{ L}+left[1-mathrm{s }/ left(mathrm{s}+upgamma right)right]*{mathrm{e}}_{mathrm{s}}*mathrm{f }left({mathrm{v}}_{mathrm{u}}right)$$with γ: psychrometric constant in hPa K−1; s: the slope of the saturation vapour pressure curve in hPa K−1; rs: short wave radiation balance in W m−2; rL: long-wave radiation balance in W m−2; L: specific evaporation heat in W m−2 mm−1 d; es: saturation deficit in hPa; f (vu): ventilation function with vu being the daily average wind speed in m s−1.Within the module cooling, the energy needed for the transition of water from liquid to gaseous phase was calculated based on the crown area (CA) and the transpiration eta sum:$${mathrm{E}}_{mathrm{A}}= {mathrm{et}}_{mathrm{a}}*mathrm{CA}-left({mathrm{L}}_{mathrm{O}}* -0.00242*mathrm{temp}right) / {mathrm{f}}_{mathrm{con}}$$with EA: energy released by a tree through transpiration (kWh tree-1), LO: energy needed for the transition of the 1 kg of water from the liquid to gaseous phase = 2.498 MJ (kgH2O)-1 and temp = temperature in ℃, fcon: 0.5.The calculation of new assimilation in the module of photosynthesis and respiration was on the basis of the approach of Haxeltine and Prenticem25. The model assumed that 50% of the incoming short-wave radiation is photosynthetic active radiation (PAR). Using the LAI and a light extinction factor of 0.5, the radiation amount of 1 m2 leaf area can be estimated based on an exponential function according to the Lambert–Beer law. This way, the gross assimilation per m2 leaf area as the daily mean of the month can be derived from:$${text{A}} = {text{d}}*{{left[ {left( {{text{J}}_{{text{p}}} + {text{J}}_{{text{r}}} – {text{sqrt}} left( {left( {{text{J}}_{{text{P}}} + {text{J}}_{{text{r}}} } right)^{2} – 4*uptheta *{text{J}}_{{text{p}}} *{text{J}}_{{text{r}}} } right)} right)} right]} mathord{left/ {vphantom {{left[ {left( {{text{J}}_{{text{p}}} + {text{J}}_{{text{r}}} – {text{sqrt}} left( {left( {{text{J}}_{{text{P}}} + {text{J}}_{{text{r}}} } right)^{2} – 4*uptheta *{text{J}}_{{text{p}}} *{text{J}}_{{text{r}}} } right)} right)} right]} {left( {2*uptheta } right)}}} right. kern-0pt} {left( {2*uptheta } right)}}$$with A: gross assimilation [g C m−2 d−1]; d: mean day length of the month [h]; Jp: reaction of photosynthesis on absorbed photosynthetic radiation [g C m−2 h−1]; Jr: rubisco limited rate of photosynthesis [g C m−2 h−1]; θ: form factor = 0.7.Jp was defined as a function of the photosynthetic active radiation PAR in mol m−2 h−1 and the efficiency of carbon fixation per absorbed PAR [g C mol−1].$${text{J}}_{{text{p}}} = {text{c}}_{{text{p}}} {text{*PAR}}$$$${text{c}}_{{text{p}}} = alpha *left( {{text{p}}_{{{text{ci}}}} – {text{r}}} right){ /}left( {{text{p}}_{{{text{ci}}}} – {text{r}}} right)*gamma *{text{m}}_{{{text{co}}_{2} }} *{text{i}}left[ {text{t}} right]$$with α: intrinsic quantum efficiency for CO2 uptake = 0.08; pci: partial pressure of the internal CO2 [Pa]; r: CO2 compensation point [Pa]; ϒ: species dependent adjustment function for tree age; m CO2: molecular mass of C = 12.0 g mol−1; i[t]: influence of temperature on efficiency.Net assimilation AN [g C m−2 d−1] was then derived from the gross assimilation A and the dark respiration Rd by:$${text{A}}_{{text{N}}} = {text{A}} – {text{R}}_{{text{d}}}$$$${text{R}}_{{text{d}}} =upbeta *{text{V}}_{{text{m}}}$$where Vm was calculated as:$${text{V}}_{{text{m}}} = {1 mathord{left/ {vphantom {1 upbeta }} right. kern-0pt} upbeta } * {{{text{c}}_{{text{p}}} } mathord{left/ {vphantom {{{text{c}}_{{text{p}}} } {{text{c}}_{{text{r}}} * {text{PAR}} * left[ {left( {2uptheta – 1} right) * beta * {{text{d}} mathord{left/ {vphantom {{text{d}} {{text{d}}_{max } }}} right. kern-0pt} {{text{d}}_{max } }} – left( {2uptheta *upbeta *{{text{d}} mathord{left/ {vphantom {{text{d}} {{text{d}}_{max } }}} right. kern-0pt} {{text{d}}_{max } }} – {text{c}}_{{text{r}}} } right)*varsigma } right]}}} right. kern-0pt} {{text{c}}_{{text{r}}} * {text{PAR}} * left[ {left( {2theta – 1} right) * beta * {{text{d}} mathord{left/ {vphantom {{text{d}} {{text{d}}_{max } }}} right. kern-0pt} {{text{d}}_{max } }} – left( {2theta *upbeta *{{text{d}} mathord{left/ {vphantom {{text{d}} {{text{d}}_{max } }}} right. kern-0pt} {{text{d}}_{max } }} – {text{c}}_{{text{r}}} } right)*varsigma } right]}}$$By multiplying AN, the number of days and the total leaf area, the entire monthly net assimilation of the tree can be obtained. In this study, we assumed a fixed share of 50% as respiration based on the gross primary production that the resulting net primary production NPP was transformed in the content of fixed carbon by multiplying the value with the carbon conversion factor 0.524.$${mathrm{Carbon}}_{mathrm{fix}}=0.5*mathrm{NPP}$$Statistical analysesThe software package R was used for statistical analysis. To investigate the differences between means, two-sampled t-test and analysis of variance (ANOVA) with Tukey’s HSD (honestly significant difference) test were used. All the cases, the means were reported as significant when P More
