LsGRP1, a class II glycine-rich protein of Lilium, confers plant resistance via mediating innate immune activation and inducing fungal programmed cell death.

Defence-related LsGRP1 is a leaf-specific plant class II glycine-rich protein (GRP) involved in salicylic acid-induced systemic resistance against grey mould caused by necrotrophic Botrytis elliptica in lily (Lilium) cultivar Stargazer. The C-terminal region of LsGRP1 (LsGRP1C ) can inhibit fungal growth in vitro via a mechanism of inducing fungal apoptosis programmed cell death (PCD). In this study, the role of LsGRP1 in induced defence mechanism was investigated using LsGRP1-silenced Stargazer lily and LsGRP1-transgenic Arabidopsis thaliana. LsGRP1 silencing in lily was found to slightly inhibit plant growth and greatly increase the susceptibility to B. elliptica by suppressing callose deposition and early reactive oxygen species (ROS) accumulation. In contrast, LsGRP1-transgenic Arabidopsis showed higher resistance to Botrytis cinerea and also to Pseudomonas syringae pv. tomato DC3000 as compared to the wild type, accompanied with the enhancement of callose deposition and ROS accumulation. Additionally, LsGRP1 silencing increased plant cell death caused by B. elliptica secretion and reduced pathogen-associated molecular pattern (PAMP)-triggered defence activation in Stargazer lily. Consistently, LsGRP1 expression boosted PAMP-triggered defence responses and effector recognition-induced hypersensitive response in Arabidopsis. Moreover, fungal apoptosis PCD triggered by LsGRP1 in an LsGRP1C -dependent manner was demonstrated by leaf infiltration with LsGRP1C -containing recombinant proteins in Stargazer lily. Based on these results, we presume that LsGRP1 plays roles in plant defence via functioning as a pathogen-inducible switch for plant innate immune activation and acting as a fungal apoptosis PCD inducer to combat pathogen attack.

Dynamic responses of DOC and DIC transport to different flow regimes in a subtropical small mountainous river.

Transport of riverine dissolved carbon (including DOC and DIC) is a crucial process linking terrestrial and aquatic C reservoirs, but has rarely been examined in subtropical small mountainous rivers (SMRs). This study monitored DOC and DIC concentrations on a biweekly basis during non-event flow periods and at 3 h intervals during two typhoon events in three SMRs in southwestern Taiwan between January 2014 and August 2016. Two models, HBV (the Hydrologiska Byråns Vattenbalansavdelning model) and a three-endmember mixing model, were applied to determine the quantities of DOC and DIC transport from different flow paths. The results show that the annual DOC and DIC fluxes were 2.7-4.8 and 48.4-54.3 t C km-2 yr-1, respectively, which were approx. 2 and 20 times higher than the global mean of 1.4 and 2.6 t C km-2 yr-1, respectively. The DIC / DOC ratio was 14.08, which is much higher than the mean of large rivers worldwide (1.86), and indicates the high rates of chemical weathering in this region. The two typhoons contributed 12%-14% of the annual streamflow in only 3 days (about 1.0% of the annual time), whereas 15.0%-23.5% and 9.2%-12.6% of the annual DOC and DIC flux, respectively, suggested that typhoons play a more important role in DOC transport than DIC transport. The end-member mixing model suggested that DOC and DIC export was mainly from surface runoff and deep groundwater, respectively. The unique patterns seen in Taiwan SMRs characterized by high dissolved carbon flux, high DIC / DOC ratio, and large transport by intense storms should be taken into consideration when estimating global carbon budgets.

Unusual Roles of Discharge, Slope and SOC in DOC Transport in Small Mountainous Rivers, Taiwan.

Riverine dissolved organic carbon (DOC), responsible for riverine productivity, is rarely documented in subtropical small mountainous rivers (SMRs) where high rainfall and steep slopes are the main features. This study investigated the DOC export at eight sites in three Taiwan SMRs to characterize the dynamics and controlling factors of DOC transport. Results showed that the mean DOC concentration of ~0.78 mg L-1 is much lower than the global average of ~5.29 mg L-1. However, the mean DOC yield, ~22.51 kg-C ha-1 yr-1, is higher than the global average of 14.4-19.3 kg-C ha-1 yr-1. Comparing with worldwide rivers from literature, the annual discharge, slope, and SOC (soil organic carbon) are controlling factors as expected, though they influence in different ways. SOC stock likely regulated by elevation-dependent biomes dominate the DOC supply, while slope restrains the DOC generation due to shallow soil depth and fast runoff velocity. However, the abundant discharge flushing this persistent low supply leads to a large DOC export in the SMRs. Furthermore, the DOC dynamics during typhoon periods showed a clockwise hysteresis, suggesting that the DOC is mainly from the riparian zone or downslope area during the rising limb of the hydrograph. This study elucidates the DOC transport in SMRs and provides an atypical yet significant piece of understanding on DOC transport in a global context.

Differences in N loading affect DOM dynamics during typhoon events in a forested mountainous catchment.

The dissolved organic matter (DOM) and nutrient dynamics in small mountainous rivers (SMRs) strongly depend on hydrologic conditions, and especially on extreme events. Here, we investigated the quantity and quality of DOM and inorganic nutrients during base-flow and typhoon events, in a chronically N-saturated mainstream and low N-loaded tributaries of a forested small mountainous reservoir catchment in Taiwan. Our results suggest that divergent transport mechanisms were triggered in the mainstream vs. tributaries during typhoons. The mainstream DON increased from 3.4 to 34.7% of the TDN pool with a static DOC:NO3-N ratio and enhanced DOM freshness, signalling a N-enriched DOM transport. Conversely, DON decreased from 46 to 6% of the TDN pool in the tributaries and was coupled with a rapid increase of the DOC:NO3-N ratio and humified DOM signals, suggesting the DON and DOC were passively and simultaneously transported. This study confirmed hydrology and spatial dimensions being the main drivers shaping the composition and concentration of DOM and inorganic nutrients in small mountainous catchments subject to hydrologic extremes. We highlighted that the dominant flow paths largely controlled the N-saturation status and DOM composition within each sub-catchment, the effect of land-use could therefore be obscured. Furthermore, N-saturation status and DOM composition are not only a result of hydrologic dynamics, but potential agents modifying the transport mechanism of solutes export from fluvial systems. We emphasize the importance of viewing elemental dynamics from the perspective of a terrestrial-aquatic continuum; and of taking hydrologic phases and individual catchment characteristics into account in water quality management.

The sources of streamwater to small mountainous rivers in Taiwan during typhoon and non-typhoon seasons.

The dynamics and behaviors of streamwater chemistry are rarely documented for subtropical small mountainous rivers. A 1-year detailed time series of streamwater chemistry, using non-typhoon and typhoon samples, was monitored in two watersheds, with and without cultivation, in central Taiwan. Rainwater, soil leachate, and well water were supplemented to explain the streamwater chemistry. The concentrations of fluoride, chloride, sulfate, magnesium, potassium, calcium, strontium, silicon, and barium of all the water samples were measured. Principal component analysis and residual analysis were applied to examine the mechanisms of solute transport and investigate possible sources contributing to the streamwater chemistry. In addition to the influence of well water and soil leachate on streamwater chemistry during non-typhoon period, overland flow and surface erosion affect streamwater chemistry during the typhoon period. The latter has not been discussed in previous studies. Surface erosion is likely to be an end member and non-conservatively mixed with other end members, resulting in a previously unobserved blank zone in the mixing space. This has a particularly great impact on small mountainous watersheds, which suffer from rapid erosion. Moreover, fertilizer contaminates agricultural soil, making soil water end members more identifiable. To our knowledge, this study is the first to clearly illustrate the dynamics and sources of streamwater chemistry of small mountainous rivers that are analogous to rivers in Oceania.

Effects of different N sources on riverine DIN export and retention in a subtropical high-standing island, Taiwan.

Increases in nitrogen (N) availability and mobility resulting from anthropogenic activities have substantially altered the N cycle, both locally and globally. Taiwan characterized by the subtropical montane landscape with abundant rainfall, downwind of the most rapidly industrializing eastern coast of China, can be a demonstration site for extremely high N input and riverine DIN (dissolved inorganic N) export. We used 49 watersheds with similar climatic and landscape settings but classified into low, moderate, and highly disturbed categories based on population density to illustrate their differences in nitrogen inputs (through atmospheric N deposition, synthetic fertilizers, and human emission) and DIN export ratios. Our results showed that the island-wide average riverine DIN export is ~ 3800 kg N km-2 yr-1, approximately 18 times the global average. The average riverine DIN export ratios are 0.30-0.51, which are much higher than the averages of 0.20-0.25 of large rivers around the world, indicating excessive N input relative to ecosystem demand or retention capacity. The low disturbed watersheds have a high N retention capacity and DIN export ratios of 0.06-0.18 in spite of the high N input (~ 4900 kg N km-2 yr-1). The high retention capacity is likely due to effective uptake by secondary forests in the watersheds. The moderately disturbed watersheds show a linear increase in DIN export with increases in total N inputs and mean DIN export ratios of 0.20 to 0.31. The main difference in land use between low and moderately disturbed watersheds is the greater proportion of agricultural land cover in the moderately disturbed watersheds. Thus, their greater DIN export could be attributed to N fertilizers used in the agricultural lands. The greater export ratios also imply that agricultural lands have a lower proportional N retention capacity and that reforestation could be an effective land management practice to reduce riverine DIN export. The export ratios of the highly disturbed watersheds are very high, 0.42-0.53, suggesting that much of the N input is transported downstream directly, and urges the need to increase the proportion of households connected to a sewage system and improve the effectiveness of wastewater treatment systems. The increases in the riverine DIN export ratio along the gradient of human disturbance also suggest a gradient in N saturation in subtropical Taiwan. Our results help to improve our understanding of factors controlling riverine DIN export and provide empirical evidence that calls for sound N emission/pollution control measures.

Apportioning riverine DIN load to export coefficients of land uses in an urbanized watershed.

The apportionment of riverine dissolved inorganic nitrogen (DIN) load to individual land use on a watershed scale demands the support of accurate DIN load estimation and differentiation of point and non-point sources, but both of them are rarely quantitatively determined in small montane watersheds. We introduced the Danshui River watershed of Taiwan, a mountainous urbanized watershed, to determine the export coefficients via a reverse Monte Carlo approach from riverine DIN load. The results showed that the dynamics of N fluctuation determines the load estimation method and sampling frequency. On a monthly sampling frequency basis, the average load estimation of the methods (GM, FW, and LI) outperformed that of individual method. Export coefficient analysis showed that the forest DIN yield of 521.5kg-Nkm(-2)yr(-1) was ~2.7-fold higher than the global riverine DIN yield (mainly from temperate large rivers with various land use compositions). Such a high yield was attributable to high rainfall and atmospheric N deposition. The export coefficient of agriculture was disproportionately larger than forest suggesting that a small replacement of forest to agriculture could lead to considerable change of DIN load. The analysis of differentiation between point and non-point sources showed that the untreated wastewater (non-point source), accounting for ~93% of the total human-associated wastewater, resulted in a high export coefficient of urban. The inclusion of the treated and untreated wastewater completes the N budget of wastewater. The export coefficient approach serves well to assess the riverine DIN load and to improve the understanding of N cascade.