Functional composition drives ecosystem function through multiple mechanisms in a broadleaved subtropical forest.

Understanding the role of biodiversity (B) in maintaining ecosystem function (EF) is a foundational scientific goal with applications for resource management and conservation. Two main hypotheses have emerged that address B-EF relationships: niche complementarity (NC) and the mass-ratio (MR) effect. We tested the relative importance of these hypotheses in a subtropical old-growth forest on the island nation of Taiwan for two EFs: aboveground biomass (ABG) and coarse woody productivity (CWP). Functional dispersion (FDis) of eight plant functional traits was used to evaluate complementarity of resource use. Under the NC hypothesis, EF will be positively correlated with FDis. Under the MR hypothesis, EF will be negatively correlated with FDis and will be significantly influenced by community-weighted mean (CWM) trait values. We used path analysis to assess how these two processes (NC and MR) directly influence EF and may contribute indirectly to EF via their influence on canopy packing (stem density). Our results indicate that decreasing functional diversity and a significant influence of CWM traits were linked to increasing AGB for all eight traits in this forest supporting the MR hypothesis. Interestingly, CWP was primarily influenced by NC and MR indirectly via their influence on canopy packing. Maximum height explained more of the variation in both AGB and CWP than any of the other plant functional traits. Together, our results suggest that multiple mechanisms operate simultaneously to influence EF, and understanding their relative importance will help to elucidate the role of biodiversity in maintaining ecosystem function.

Quantifying effects of habitat heterogeneity and other clustering processes on spatial distributions of tree species.

Spatially explicit consideration of species distribution can significantly add to our understanding of species coexistence. In this paper, we evaluated the relative importance of habitat heterogeneity and other clustering processes (e.g., dispersal limitation, collectively called the non-habitat clustering process) in explaining the spatial distribution patterns of 341 tree species in three stem-mapped 25-50 ha plots of tropical, subtropical, and temperate forests. Their relative importance was estimated by a method that can take one mechanism into account when estimating the effects of the other mechanism and vice versa. Our results demonstrated that habitat heterogeneity was less important in explaining the observed species patterns than other clustering processes in plots with flat topography but was more important in one of the three plots that had a complex topography. Meanwhile, both types of clustering mechanisms (habitat or non-habitat) were pervasive among species at the 50-ha scale across the studied plots. Our analyses also revealed considerable variation among species in the relative importance of the two types of mechanism within each plot and showed that this species-level variation can be partially explained by differences in dispersal mode and growth form of species in a highly heterogeneous environment. Our findings provide new perspectives on the formation of species clustering. One important finding is that a significant species-habitat association does not necessarily mean that the habitat heterogeneity has a decisive influence on species distribution. The second insight is that the large species-level variation in the relative importance of the two types of clustering mechanisms should not be ignored. Non-habitat clustering processes can play an important role on species distribution.

Chelator effects on bioconcentration and translocation of cadmium by hyperaccumulators, Tagetes patula and Impatiens walleriana.

French marigold (Tagetes patula) and impatiens (Impatiens walleriana) can act as hyperaccumulator plants for removal of cadmium (Cd) from contaminated sites. In this study, an exponential decay model was used to predict the maximum removal of Cd from artificially spiked soils by impatiens. Application of a chelator, EDTA, was also assessed for effects on the bioconcentration (BCF) and translocation (TF) factors of the two species with four replicates. Exposure to Cd significantly decreased the biomass of two plant species. Impatiens and French marigold accumulated Cd at a rate of 200-1200 mg Cd kg(-1) in shoots, with BCFs and TFs of 8.5-15 and 1.7-2.6, respectively.

Quantifying nickel in soils and plants in an ultramafic area in Philippines.

In this study, concentrations of nickel (Ni) were quantified in the soils and plants in the agricultural areas of Salcedo watershed in Eastern Samar Island, Philippines. The quantity of total Ni in soils (TS-Ni) was significantly high with a mean of 1,409 mg kg(-1), while the soil available Ni (SA-Ni) was low with a mean of 8.66 mg kg(-1). As the levels of TS-Ni in the Salcedo watershed greatly exceeded the maximum allowable concentrations for agricultural soils, the site is not suitable for agricultural purposes. Despite significant TS-Ni levels, SA-Ni levels were very low due to tight binding between Ni and soil components. Consequently, all plants investigated did not meet the criterion for a Ni hyperaccumulator plant with low Ni contents (mean TP-Ni of 14.7 mg kg(-1)). Comparison of Ni levels between food plants and its recommended daily intake (RDI) suggests that consumption of food-plants grown in the study area is unlikely to pose health risks. However, caution must be taken against combined consumption of food plants with high Ni levels or their prolonged consumption, as it can induce accumulation of Ni above RDI.

The influence of EDTA application on the interactions of cadmium, zinc, and lead and their uptake of rainbow pink (Dianthus chinensis).

Soil used in this study was artificially contaminated with Cd, Zn, Pb, or applied in combinations (Cd-Zn, Cd-Pb, Zn-Pb, or Cd-Zn-Pb) to study the interactions of metals in soil contaminated with multiple metals. After planting rainbow pink (Dianthus chinensis) in these soils for 21 days, three different concentrations of ethylenedinitrilotetraacetic acid (EDTA) solutions were added to study the effect of applying EDTA on the interactions among these metals. The concentrations of Cd, Zn, and Pb in the soil solutions of different metals-treated soils increased significantly after applying 5 mmol EDTA kg(-1) soil (p<0.05). The potential of groundwater contamination will increase after applying EDTA and it is not recommended to be in situ used or have to use very carefully. The existence of Pb in the Cd-contaminated soil enhanced the uptake of Cd in rainbow pink in the treatments of control and 2 mmol EDTA kg(-1) soil. Cadmium inhibited the concentration of Zn without applying EDTA. However, whether the application of EDTA or not and the applied EDTA concentration had the greatest effect on the uptake of Pb when compared to Cd and Zn. After applying 5 mmol EDTA kg(-1) soil, Cd or Zn in the Pb-contaminated soil inhibited the uptake of Pb in rainbow pink, but there were no effect in other treatments.

Linking leaf veins to growth and mortality rates: an example from a subtropical tree community.

A fundamental goal in ecology is to link variation in species function to performance, but functional trait-performance investigations have had mixed success. This indicates that less commonly measured functional traits may more clearly elucidate trait-performance relationships. Despite the potential importance of leaf vein traits, which are expected to be related to resource delivery rates and photosynthetic capacity, there are few studies, which examine associations between these traits and demographic performance in communities. Here, we examined the associations between species traits including leaf venation traits and demographic rates (Relative Growth Rate, RGR and mortality) as well as the spatial distributions of traits along soil environment for 54 co-occurring species in a subtropical forest. Size-related changes in demographic rates were estimated using a hierarchical Bayesian approach. Next, Kendall's rank correlations were quantified between traits and estimated demographic rates at a given size and between traits and species-average soil environment. Species with denser venation, smaller areoles, less succulent, or thinner leaves showed higher RGR for a wide range of size classes. Species with leaves of denser veins, larger area, cheaper construction costs or thinner, or low-density wood were associated with high mortality rates only in small size classes. Lastly, contrary to our expectations, acquisitive traits were not related to resource-rich edaphic conditions. This study shows that leaf vein traits are weakly, but significantly related to tree demographic performance together with other species traits. Because leaf traits associated with an acquisitive strategy such as denser venation, less succulence, and thinner leaves showed higher growth rate, but similar leaf traits were not associated with mortality, different pathways may shape species growth and survival. This study suggests that we are still not measuring some of key traits related to resource-use strategies, which dictate the demography and distributions of species.

The EDTA effect on phytoextraction of single and combined metals-contaminated soils using rainbow pink (Dianthus chinensis).

Rainbow pink (Dianthus chinensis), a potential phytoextraction plant, can accumulate high concentrations of Cd from metal-contaminated soils. The soils used in this study were artificially added with different metals including (1) CK: original soil, (2) Cd-treated soil: 10 mg Cd kg(-1), (3) Zn-treated soil: 100 mg Zn kg(-1), (4) Pb-treated soil: 1000 mg Pb kg(-1), (5) Cd-Zn-treated soil: 10 mg Cd kg(-1) and 100 mg Zn kg(-1), (6) Cd-Pb-treated soil: 10 mg Cd kg(-1) and 1000 mg Pb kg(-1), (7) Zn-Pb-treated soil: 100 mg Zn kg(-1) and 1000 mg Pb kg(-1), and (8) Cd-Zn-Pb-treated soil: 10 mg Cd kg(-1), 100 mg Zn kg(-1), and 1000 mg Pb kg(-1). Three concentrations of 2Na-EDTA solutions (0 (control), 2, and 5 mmol kg(-1) soil) were added to the different metals-treated soils to study the influence of applied EDTA on single and combined metals-contaminated soils phytoextraction using rainbow pink. The results showed that the Cd, Zn, Pb, Fe, or Mn concentrations in different metals-treated soil solutions significantly increased after applying 5 mmol EDTA kg(-1) (p<0.05). The metal concentrations in different metals-treated soils extracted by deionized water also significantly increased after applying 5 mmol EDTA kg(-1) (p<0.05). Because of the high extraction capacity of both 0.005 M DTPA (pH 5.3) and 0.05 M EDTA (pH 7.0), applying EDTA did not significantly increase the Cd, Zn, or Pb concentration in both extracts for most of the treatments. Applying EDTA solutions can significantly increase the Cd and Pb concentrations in the shoots of rainbow pink (p<0.05). However, this was not statistically significant for Zn because of the low Zn concentration added into the contaminated soils. The results from this study indicate that applying 5 mmol EDTA kg(-1) can significantly increase the Cd, Zn, or Pb concentrations both in the soil solution or extracted using deionized water in single or combined metals-contaminated soils, thus increasing the accumulated metals concentrations in rainbow pink shoots. The proposed method worked especially well for Pb (p<0.05). The application of 2 mmol EDTA kg(-1) might too low to enhance the phytoextraction effect when used in silty clay soils.

Effects of EDTA on solubility of cadmium, zinc, and lead and their uptake by rainbow pink and vetiver grass.

Rainbow pink (Dianthus chinensis), a potential phytoextraction plant, can accumulate high concentrations of Cd from contaminated soils. Vetiver grass (Vetiver zizanioides) has strong and long root tissues and is a potential phytostabilization plant since it can tolerate and grow well in soils contaminated with multiple heavy metals. Soil was moderately artificially contaminated by cadmium (20 mg/kg), zinc (500 mg/kg), and lead (1000 mg/kg) in pot experiments. Three concentrations of Na2-EDTA solution (0, 5, and 10 mmol/kg soil) were added to the contaminated soils to study the influence of EDTA solution on phytoextraction by rainbow pink or phytostabilization by vetiver grass. The results showed that the concentrations of Cd, Zn, and Pb in a soil solution of rainbow pink significantly increased following the addition of EDTA (p < 0.05). The concentrations of Cd and Pb in the shoots of rainbow pink also significantly increased after EDTA solution was applied (p < 0.05), but the increase for Zn was insignificant. EDTA treatment significantly increased the total uptake of Pb in the shoot, over that obtained with the control treatment (p < 0.001), but it did not significantly increase the total uptake of Cd and Zn. The concentrations of Zn and Pb in the shoots of rainbow pink are significantly correlated with those in the soil solution, but no relationship exists with concentrations in vetiver grass. The toxicity of highly contaminating metals did not affect the growth of vetiver grass, which was found to grow very well in this study. Results of this study indicate that rainbow pink can be considered to be a potential phytoextraction plant for removing Cd or Zn from metal-contaminated soils, and that vetiver grass can be regarded as a potential phytostabilization plant that can be grown in a site contaminated with multiple heavy metals.

Effect of chemical amendments on the concentration of cadmium and lead in long-term contaminated soils.

The availability of metal in contaminated soil can be reduced by the addition of soil amendments. The objectives of this study are to study the effects of applying different soil amendments on the concentration of Cd and Pb in soil solution, DTPA or EDTA extractable Cd and Pb, and the uptake of Cd and Pb by wheat (Triticum vulgare) when growing in long-term Cd and Pb-contaminated soils, more than 20 years. The soil amendments, including check, compost, zinc oxide, calcium carbonate, calcium carbonate mixed with zinc oxide, and calcium carbonate mixed with compost, were conducted in a four replicates pot cultural study. The amended soils were incubated for six months under 60% of water holding capacity. Following incubation, wheat was grown for four months in greenhouse. Analyses of Cd concentration demonstrated a significant decrease in soil solution concentration and DTPA or EDTA extractable in soils amended with calcium carbonate or calcium carbonate mixed with ZnO (or compost) (p<0.01). These amendments can significantly reduce the Cd concentration in the grain, leaf and stem, or reduce the total Cd uptake in all parts of wheat species grown in highly contaminated soil amended with calcium carbonate or calcium carbonate mixed with ZnO (or compost) (p<0.01). The concentration of Cd in soil solution and extracted with DTPA or EDTA can predict the Cd concentration in wheat, especially for soil solution.

Food safety and bioavailability evaluations of four vegetables grown in the highly arsenic-contaminated soils on the Guandu Plain of northern Taiwan.

Arsenic contamination in a large area of agricultural fields on the Guandu Plain of northern Taiwan was confirmed in a survey conducted in 2006, but research concerning the relationship between bioavailable As concentrations in contaminated soils and crop production in Taiwan is not available. Pot experiments were conducted to examine the growth and accumulation of As in four vegetable crops grown in As-contaminated soils and to assess As intake through consumption. The phytotoxic effects of As in soils were not shown in the pot experiments in which vegetable crops were grown in soils contaminated with different As levels in situ collected from Guandu Plain (120-460 mg/kg) or artificially spiked As-contaminated soils (50-170 mg/kg). Experimental results showed that the bioavailable As extracted with 0.5M NaHCO3 from soils can be used to estimate As concentrations in vegetables. The As concentrations in the vegetables were compared with data shown in the literature and As limits calculated from drinking water standards and the provisional tolerance weekly intake (PTWI) of inorganic As established by the Food and Agriculture Organization of the United Nations/World Health Organization (FAO/WHO). Although the As levels in the vegetables were not high and the bioavailability of As in the soils was quite low, long-term consumption may result in higher As intake in the human body.

Cadmium-induced physiological response and antioxidant enzyme changes in the novel cadmium accumulator, Tagetes patula.

The accumulation and effect of cadmium (Cd) on the growth and enzymatic activities changes of antioxidants in Tagetes patula, French marigold, were investigated to reveal the physiological mechanisms corresponding to its Cd tolerance and accumulation. Hydroponically grown T. patula plants were treated with different concentrations of Cd (0, 10, 25, 50 µM Cd Cl(2)) at various regime of times. T. patula accumulated Cd to a maximum of 450 mg Cd kg(-1) dry weight (DW) in shoot and 3500 mg Cd kg(-1) DW in root after 14 days' exposure at 10 and 50 µM Cd Cl(2), respectively. The translocation factors of Cd were greater than 1 in plants exposed to 10 µM Cd Cl(2). Toxic effects were gradually observed with increasing Cd concentration (25 and 50 µM) accompanied with the reduction of biomass, chlorophyll content, decrease of cell viability and the increase level of lipid peroxidation. In leaves of T. patula, the activities of ascorbate peroxidase (APX), glutathione reductase (GR) and superoxide dismutase (SOD) were induced by Cd. However, in roots, activities of APX, GR, SOD and catalase (CAT) were significantly reduced by 25 and 50 µM Cd treatment but not 10 µM Cd. In-gel zymography analysis revealed that Cd induced the enzymatic activities of APX, MnSOD, CuZnSOD and different isozymes of GR in leaves. These results indicate that T. patula is a novel Cd accumulator and able to tolerate with Cd-induced toxicity by activation of its antioxidative defense system.

Bioavailability assessment and accumulation by five garden flower species grown in artificially cadmium-contaminated soils.

Many studies have been conducted on phytoextraction; however, non-native hyperaccumulator species are not suitable for the natural environment of Taiwan in many cases. Drawing upon previous results, the growth and heavy metal accumulation in artificially cadmium-contaminated soils were compared for five local garden flower species. The treatments included a control (CK), 9.73 +/- 0.05 mg kg(-1) (Cd-10), and 17.6 +/- 0.8 mg kg(-1) (Cd-20). All plants were harvested at 35 days after transplanting and analyzed for Cd content. Cd accumulation in the shoot of French marigold (Tagetes patula L.) and Impatiens (Impatiens walleriana Hook. f.) grown in Cd-20 treatment were 66.3 +/- 6.5 and 100 +/- 11 mg kg(-1), which equated to a removal of 0.80 +/- 0.11 and 0.60 +/- 0.37 mg Cd plant(-1), respectively. The maximum Cd accumulation of Impatiens reached the threshold value (100 mg kg(-1)) characteristic of a Cd hyperaccumulator and its bioconcentration factor (BCF) and translocation factor (TF) were greater than one. Impatiens therefore has the potential to hyperaccumulate Cd from Cd-contaminated soils. With the exception of Garden verbena, significant relationships were found between Cd concentrations in soil extracted by 0.05 M EDTA, 0.005 M DTPA, and 0.01 M CaCl2 and the concentration of Cd in the shoots of the tested garden flowers.

Large-area experiment on uptake of metals by twelve plants growing in soils contaminated with multiple metals.

A site in central Taiwan with an area of 1.3 ha and contaminated with Cr, Cu, Ni, and Zn was selected to examine the feasibility of phytoextraction. Based on the results of a preexperiment at this site, a total of approximately 20,000 plants of 12 species were selected from plants of 33 tested species to be used in a large-area phytoextraction experiment at this site. A comparison with the initial metal concentration of 12 plant species before planting demonstrated that most species accumulated significant amounts of Cr, Cu, Ni, and Zn in their shoots after growing in this contaminated site for 31 d. Among the 12 plant species, the following accumulated higher concentrations of metals in their shoots; Garden canna and Garden verbena (45-60 mg Cr kg(-1)), Chinese ixora and Kalanchoe (30 mg Cu kg(-1)), Rainbow pink and Sunflower (30 mg Ni kg(-1)), French marigold and Sunflower (300-470 mg Zn kg(-1)). The roots of the plants of most of the 12 plant species can accumulate higher concentrations of metals than the shoots and extending the growth period promotes accumulation in the shoots. Large-area experiments demonstrated that phytoextraction is a feasible method to enable metal-contaminated soil in central Taiwan to be reused.

Health risk-based assessment and management of heavy metals-contaminated soil sites in Taiwan.

Risk-based assessment is a way to evaluate the potential hazards of contaminated sites and is based on considering linkages between pollution sources, pathways, and receptors. These linkages can be broken by source reduction, pathway management, and modifying exposure of the receptors. In Taiwan, the Soil and Groundwater Pollution Remediation Act (SGWPR Act) uses one target regulation to evaluate the contamination status of soil and groundwater pollution. More than 600 sites contaminated with heavy metals (HMs) have been remediated and the costs of this process are always high. Besides using soil remediation techniques to remove contaminants from these sites, the selection of possible remediation methods to obtain rapid risk reduction is permissible and of increasing interest. This paper discusses previous soil remediation techniques applied to different sites in Taiwan and also clarified the differences of risk assessment before and after soil remediation obtained by applying different risk assessment models. This paper also includes many case studies on: (1) food safety risk assessment for brown rice growing in a HMs-contaminated site; (2) a tiered approach to health risk assessment for a contaminated site; (3) risk assessment for phytoremediation techniques applied in HMs-contaminated sites; and (4) soil remediation cost analysis for contaminated sites in Taiwan.

Pot experiment to study the uptake of Cd and Pb by three Indian mustards (Brassica juncea) grown in artificially contaminated soils.

Most of the metals-contaminated and fallow lands in Taiwan are a result of irrigation with illegal effluent of factories. Phytoextraction methods can be applied to reach the target of fallow-lands reuse and earn more incomes for farmers. In many studies, Indian mustards (Brassica juncea) were planted in the metal-contaminated soils to study their suitability in phytoextraction. However, the total removal of metals by plants was quite different between accessions. In this pot study, three accessions of B. juncea (cv. 182921, cv. 211000, and cv. 426308) were planted in artificially Cd- or Pb-contaminated soils to investigate the differences between them. EDTA was applied to study its effect in increasing the bioavailability of Cd and Pb and their uptake by these Indian mustards. Experimental result showed that three accessions of Indian mustard can accumulate a high concentration of Cd and Pb when growing in the artificially Cd- and Pb-contaminated soils. Their shoot Cd or Pb concentrations were significantly enhanced, resulting from the application of EDTA. Among the three accessions, B. juncea cv. 211000 accumulated the highest concentrations of Cd and Pb in their shoots compared with B. juncea cv. 182921 and cv. 426308, but its total removal was the lowest due to its lower biomass.

Baseline concentrations of ten metals in the freshwater sediments of a watershed in Taiwan.

Freshwater sediments and water samples were quarterly collected at different sampling stations from August 1996 to April 1998 in the Fei-Tsui reservoir watershed (FTRW) of northern Taiwan. The sediment and soil samples were digested using mixture of inorganic acids to analyze the total concentration of ten metals. The baseline concentrations of ten metals in the sediments of FTRW were calculated from the geometric mean (GM) values and the geometric standard deviation (GSD). The baseline concentrations of 10 metals in the 239 sediments of this watershed are listed as followings (mg/kg, only % for Fe): As 1.80-51.5, Cd 0.04-7.54, Cr 5.25-273, Cu 2.65-91.7, Hg 0.03-0.26, Mn 18.6-2310, Ni 3.95-167, Pb 1.10-75.4, Zn 6.46-327, and Fe 0.47-9.63%. However, the GMs of 10 metal concentrations in the sediments were lower than the regulated threshold concentrations in Taiwan rural soils. Highly correlations among different metals in the sediments of FTRW indicate that very similar sediments of formation process control the metal content associated with the parent materials. Inadequate land use results in As, Cr, Mn, and Ni accumulated in the sediment to show potentially contamination risk in the study area associated with accelerated erosion and runoff.