Messages from the other side: parasites receive damage cues from their host plants

As sessile organisms, plants rely on their environment for cues indicating imminent herbivory. These cues can originate from tissues on the same plant or from different individuals. Since parasitic plants form vascular connections with their host, parasites have the potential to receive cues from hosts that allow them to adjust defenses against future herbivory. However, the role of plant communication between hosts and parasites for herbivore defense remains poorly investigated. Here we examined the effects of damage to lupine hosts (Lupinus texensis) on responses of the attached hemiparasite (Castilleja indivisa), and indirectly, on a specialist herbivore of the parasite, buckeyes (Junonia coenia). Lupines produce alkaloids as defenses against herbivore that can be taken up by the parasite. We found that damage to lupine host plants by beet armyworm (Spodoptera exigua) significantly increased jasmonic acid (JA) levels in both the lupine host and parasite, suggesting uptake of phytohormones or priming of parasite defenses using host cues. However, lupine host damage did not induce changes in alkaloid levels in the hosts or parasites. Interestingly, the parasite had substantially higher concentrations of JA and alkaloids compared to lupine host plants. Buckeye herbivores consumed more parasite tissue when attached to damaged compared to undamaged hosts. We hypothesize that increased JA due to lupine host damage induced higher iridoid glycosides in the parasite, which are feeding stimulants for this specialist herbivore. Our results demonstrate that damage to hosts may affect both parasites and associated herbivores, indicating cascading effects of host damage on multiple trophic levels

Organic phosphorus cycling may control grassland responses to nitrogen deposition: A long-term field manipulation and modelling study

This is the final version. Available on open access from the European Geosciences Union via the DOI in this recordData availability: Data presented in the manuscript have been deposited with NERC's Environmental Information Data Centre (EIDC) at the following DOI: https://doi.org/10.5285/98b473c7-3ca9-498d-a851-31152b1f1da7 (Taylor et al., 2021). All data to be archived are present in the Supplement for review purposes.Ecosystems limited in phosphorous (P) are widespread, yet there is limited understanding of how these ecosystems may respond to anthropogenic deposition of nitrogen (N) and the interconnected effects on the biogeochemical cycling of carbon (C), N, and P. Here, we investigate the consequences of enhanced N addition for the C-N-P pools of two P-limited grasslands, one acidic and one limestone, occurring on contrasting soils, and we explore their responses to a long-term nutrient-manipulation experiment. We do this by combining data with an integrated C-N-P cycling model (N14CP). We explore the role of P-access mechanisms by allowing these to vary in the modelling framework and comparing model plant-soil C-N-P outputs to empirical data. Combinations of organic P access and inorganic P availability most closely representing empirical data were used to simulate the grasslands and quantify their temporal response to nutrient manipulation. The model suggested that access to organic P is a key determinant of grassland nutrient limitation and responses to experimental N and P manipulation. A high rate of organic P access allowed the acidic grassland to overcome N-induced P limitation, increasing biomass C input to soil and promoting soil organic carbon (SOC) sequestration in response to N addition. Conversely, poor accessibility of organic P for the limestone grassland meant N provision exacerbated P limitation and reduced biomass input to the soil, reducing soil carbon storage. Plant acquisition of organic P may therefore play an important role in reducing P limitation and determining responses to anthropogenic changes in nutrient availability. We conclude that grasslands differing in their access to organic P may respond to N deposition in contrasting ways, and where access is limited, soil organic carbon stocks could decline.Natural Environment Research Council (NERC

Transpiration from subarctic deciduous woodlands: environmental controls and contribution to ecosystem evapotranspiration

Potential land‐climate feedbacks in subarctic regions, where rapid warming is driving forest expansion into the tundra, may be mediated by differences in transpiration of different plant functional types. Here we assess the environmental controls of overstorey transpiration and its relevance for ecosystem evapotranspiration in subarctic deciduous woodlands. We measured overstorey transpiration of mountain birch canopies and ecosystem evapotranspiration in two locations in northern Fennoscandia, having dense (Abisko) and sparse (Kevo) overstories. For Kevo, we also upscale chamber‐measured understorey evapotranspiration from shrubs and lichen using a detailed land cover map. Sub‐daily evaporative fluxes were not affected by soil moisture, and showed similar controls by vapour pressure deficit and radiation across sites. At the daily timescale, increases in evaporative demand led to proportionally higher contributions of overstorey transpiration to ecosystem evapotranspiration. For the entire growing season, the overstorey transpired 33% of ecosystem evapotranspiration in Abisko and only 16% in Kevo. At this latter site, the understorey had a higher leaf area index and contributed more to ecosystem evapotranspiration compared to the overstorey birch canopy. In Abisko, growing season evapotranspiration was 27% higher than precipitation, consistent with a gradual soil moisture depletion over the summer. Our results show that overstorey canopy transpiration in subarctic deciduous woodlands is not the dominant evaporative flux. However, given the observed environmental sensitivity of evapotranspiration components, the role of deciduous trees in driving ecosystem evapotranspiration may increase with the predicted increases in tree cover and evaporative demand across subarctic regions

Nitrogen Deposition Reduces Plant Diversity and Alters Ecosystem Functioning: Field-Scale Evidence from a Nationwide Survey of UK Heathlands

Findings from nitrogen (N) manipulation studies have provided strong evidence of the detrimental impacts of elevated N deposition on the structure and functioning of heathland ecosystems. Few studies, however, have sought to establish whether experimentally observed responses are also apparent under natural, field conditions. This paper presents the findings of a nationwide field-scale evaluation of British heathlands, across broad geographical, climatic and pollution gradients. Fifty two heathlands were selected across an N deposition gradient of 5.9 to 32.4 kg ha−1 yr−1. The diversity and abundance of higher and lower plants and a suite of biogeochemical measures were evaluated in relation to climate and N deposition indices. Plant species richness declined with increasing temperature and N deposition, and the abundance of nitrophilous species increased with increasing N. Relationships were broadly similar between upland and lowland sites, with the biggest reductions in species number associated with increasing N inputs at the low end of the deposition range. Both oxidised and reduced forms of N were associated with species declines, although reduced N appears to be a stronger driver of species loss at the functional group level. Plant and soil biochemical indices were related to temperature, rainfall and N deposition. Litter C:N ratios and enzyme (phenol-oxidase and phosphomonoesterase) activities had the strongest relationships with site N inputs and appear to represent reliable field indicators of N deposition. This study provides strong, field-scale evidence of links between N deposition - in both oxidised and reduced forms - and widespread changes in the composition, diversity and functioning of British heathlands. The similarity of relationships between upland and lowland environments, across broad spatial and climatic gradients, highlights the ubiquity of relationships with N, and suggests that N deposition is contributing to biodiversity loss and changes in ecosystem functioning across European heathlands

Contrasting effects of hemiparasites on ecosystem processes: can positive litter effects offset the negative effects of parasitism?

Hemiparasites are known to influence community structure and ecosystem functioning, but the underlying mechanisms are not well studied. Variation in the impacts of hemiparasites on diversity and production could be due to the difference in the relative strength of two interacting pathways: direct negative effects of parasitism and positive effects on N availability via litter. Strong effects of parasitism should result in substantial changes in diversity and declines in productivity. Conversely, strong litter effects should result in minor changes in diversity and increased productivity. We conducted field-based surveys to determine the association of Castillejaoccidentalis with diversity and productivity in the alpine tundra. To examine litter effects, we compared the decomposition of Castilleja litter with litter of four other abundant plant species, and examined the decomposition of those four species when mixed with Castilleja. Castilleja was associated with minor changes in diversity but almost a twofold increase in productivity and greater foliar N in co-occurring species. Our decomposition trials suggest litter effects are due to both the rapid N loss of Castilleja litter and the effects of mixing Castilleja litter with co-occurring species. Castilleja produces litter that accelerates decomposition in the alpine tundra, which could accelerate the slow N cycle and boost productivity. We speculate that these positive effects of litter outweigh the effects of parasitism in nutrient-poor systems with long-lived hemiparasites. Determining the relative importance of parasitism and litter effects of this functional group is crucial to understand the strong but variable roles hemiparasites play in affecting community structure and ecosystem processes

Summer warming explains widespread but not uniform greening in the Arctic tundra biome

Arctic warming can influence tundra ecosystem function with consequences for climate feedbacks, wildlife and human communities. Yet ecological change across the Arctic tundra biome remains poorly quantified due to field measurement limitations and reliance on coarse-resolution satellite data. Here, we assess decadal changes in Arctic tundra greenness using time series from the 30 m resolution Landsat satellites. From 1985 to 2016 tundra greenness increased (greening) at ~37.3% of sampling sites and decreased (browning) at ~4.7% of sampling sites. Greening occurred most often at warm sampling sites with increased summer air temperature, soil temperature, and soil moisture, while browning occurred most often at cold sampling sites that cooled and dried. Tundra greenness was positively correlated with graminoid, shrub, and ecosystem productivity measured at field sites. Our results support the hypothesis that summer warming stimulated plant productivity across much, but not all, of the Arctic tundra biome during recent decades

Plant defences mediate interactions between herbivory and the direct foliar uptake of atmospheric reactive nitrogen

Reactive nitrogen from human sources (e.g., nitrogen dioxide, NO2) is taken up by plant roots following deposition to soils, but can also be assimilated by leaves directly from the atmosphere. Leaf uptake should alter plant metabolism and overall nitrogen balance and indirectly influence plant consumers; however, these consequences remain poorly understood. Here we show that direct foliar assimilation of NO2 increases levels of nitrogen-based defensive metabolites in leaves and reduces herbivore consumption and growth. These results suggest that atmospheric reactive nitrogen could have cascading negative effects on communities of herbivorous insects. We further show that herbivory induces a decrease in foliar uptake, indicating that consumers could limit the ability of vegetation to act as a sink for nitrogen pollutants (e.g., smog from mobile emissions). Our study suggests that the interactions of foliar uptake, plant defence and herbivory could have significant implications for understanding the environmental consequences of reactive nitrogen