Dynamics and mechanisms of secondary invasion following biological control of an invasive plant.

Secondary invasions in which nontarget invaders expand following eradication of a target invader commonly occur in habitats with multiple invasive plant species and can prevent recovery of native communities. However, the dynamics and mechanisms of secondary invasion remain unclear. Here, we conducted a common garden experiment to test underlying mechanisms of secondary invasion for 14 nontarget invaders after biological control of Ambrosia artemisiifolia in two consecutive years. We found secondary invasion for all tested nontarget invaders, but secondary invasiveness (change relative to natives) varied with species and time. Specifically, secondary invasiveness depended most strongly on phylogenetic relatedness between the target and nontarget invaders in the first year with closely related nontarget invaders being most invasive. By contrast, secondary invasiveness in the second year was mostly driven by functional traits with taller nontarget invaders or those with higher specific leaf area, or smaller seeds especially invasive. Our study indicates that secondary invasion is likely to occur wherever other invasive plants co-occur with an invasive species targeted for control. Furthermore, the most problematic invaders will initially be species closely related to the target invader but then species with rapid growth and high reproduction are most likely to be more aggressive secondary invaders.

Fluctuations in resource availability shape the competitive balance among non-native plant species.

Fluctuating resource availability plays a critical role in determining non-native plant invasions through mediating the competitive balance between non-native and native species. However, the impact of fluctuating resource availability on interactions among non-native species remains largely unknown. This represents a barrier to understanding invasion mechanisms, particularly in habitats that harbor multiple non-native species with different responses to fluctuating resource availability. To examine the responses of non-native plant species to nutrient fluctuations, we compared the growth of each of 12 non-native species found to be common in local natural areas to nutrients supplied at a constant rate or supplied as a single large pulse in a pot experiment. We found that seven species produced more biomass with pulsed nutrients compared to constant nutrients (hereafter "benefitting species"), while the other five species did not differ between nutrient enrichment treatments (hereafter "non-benefitting species"). To investigate how nutrient fluctuations influence the interactions among non-native plant species, we established experimental non-native communities in the field with two benefitting and two non-benefitting non-native species. Compared with constant nutrient supply, the single large pulse of nutrient did not influence community biomass, but strongly increased the biomass and cover of the benefitting species and decreased those of the non-benefitting species. Furthermore, the benefitting species had higher leaf N content and greater plant height when nutrients were supplied as a single large pulse than at a constant rate, whereas the non-benefitting species showed no differences in leaf N content and were shorter when nutrients were supplied as a single large pulse than at a constant rate. Our results add to the growing evidence that the individual responses of non-native species to nutrient fluctuation are species-specific. More importantly, benefitting species were favored by nutrients coming in a pulse, while non-benefitting ones were favored by nutrients coming constantly when they grew together. This suggests that nutrient fluctuations can mediate the competitive balance among non-native plants and may thus determine their invasion success in a community harboring multiple non-native plant species.

Nitrogen input enhances microbial carbon use efficiency by altering plant-microbe-mineral interactions.

Microbial growth and respiration are at the core of the soil carbon (C) cycle, as these microbial physiological performances ultimately determine the fate of soil C. Microbial C use efficiency (CUE), a critical metric to characterize the partitioning of C between microbial growth and respiration, thus controls the sign and magnitude of soil C-climate feedback. Despite its importance, the response of CUE to nitrogen (N) input and the relevant regulatory mechanisms remain poorly understood, leading to large uncertainties in predicting soil C dynamics under continuous N input. By combining a multi-level field N addition experiment with a substrate-independent 18 O-H2 O labelling approach as well as high-throughput sequencing and mineral analysis, here we elucidated how N-induced changes in plant-microbial-mineral interactions drove the responses of microbial CUE to N input. We found that microbial CUE increased significantly as a consequence of enhanced microbial growth after 6-year N addition. In contrast to the prevailing view, the elevated microbial growth and CUE were not mainly driven by the reduced stoichiometric imbalance, but strongly associated with the increased soil C accessibility from weakened mineral protection. Such attenuated organo-mineral association was further linked to the N-induced changes in the plant community and the increased oxalic acid in the soil. These findings provide empirical evidence for the tight linkage between mineral-associated C dynamics and microbial physiology, highlighting the need to disentangle the complex plant-microbe-mineral interactions to improve soil C prediction under anthropogenic N input.

The effects of plant-soil feedback on invasion resistance are soil context dependent.

There is growing interest in understanding the role that plant-soil feedbacks (PSFs) may play in invasion resistance. However, recent studies have shown that there is great uncertainty in explaining community patterns by PSF studies regarding invasions. This uncertainty may be partly because soils used for PSF studies are usually collected from open areas rather than natural communities, thus ignoring the effects of community contexts that may specifically influence the soil feedbacks of community residents to invaders. We performed a two-phase pot experiment to study the soil feedback initiated by ten co-occurring native and exotic species to a forest invader, Phytolacca americana, and the experiments were performed in forest soil and open area soil. The context-dependent mechanisms were further explored by studying different components of PSF. The results showed that natives and exotics had positive and negative effects on P. americana in the open area soil, respectively, but both had negative effects in the forest soil. Nutrient limitation was more important for the PSF in open area soil, whereas biotic factors were likely the primary mechanisms explaining the PSF in forest soil. Additionally, the litter-mediated allelopathy of dominant Quercus acutissima caused the strongest inhibition of the invader. These results suggest that native species can effectively resist invasion by producing negative PSF depending on the community context. Evidence that exotic species promote invasion through positive PSFs was not obtained. This study provided preliminary insights into the possibility of bridging PSF studies and community patterns.

Mixed evidence for plant-soil feedbacks in forest invasions.

Plant-soil feedbacks (PSFs) are plant-mediated changes to soil properties that ultimately influence plant performance, and can, thus, determine plant diversity, succession, and invasion. We hypothesized that PSFs influence invasion processes and that PSF mechanisms are largely driven by changes in soil properties produced by specific plant species. To test these hypotheses, we studied the effects of different soils collected from under common plant species on the growth of the invasive plant Phytolacca americana. We found that PSFs may interfere with invasion resistance because P. americana seedlings showed reduced growth (lower biomass) in soils collected from underneath some native species compared with soils collected from underneath P. americana and two non-native plants. We then selected eight co-occurring native and non-native plant species, and examined PSF dynamics and mechanisms in a pairwise conditioned soil greenhouse experiment. Plant species-specific conditioning effects regarding soil nutrients and enzyme activities were observed. Phytolacca americana had a high ability to use soil N, which may be related to its high invasion ability. Soil P was significantly lower in Quercus acutissima-conditioned soil, indicating that low P availability in Q. acutissima forests may enhance resistance to plant invasion. However, surprisingly, some native plants did not produce PSF effects that decreased the relative performance of invasive plants, nor did the invasive plants produce PSF effects that increased their own performance. We speculate that these PSF findings from greenhouse experiments cannot be extrapolated to field conditions because the litter and allelochemicals of some plants may be important for invasion resistance.

Regulation of priming effect by soil organic matter stability over a broad geographic scale.

The modification of soil organic matter (SOM) decomposition by plant carbon (C) input (priming effect) represents a critical biogeochemical process that controls soil C dynamics. However, the patterns and drivers of the priming effect remain hidden, especially over broad geographic scales under various climate and soil conditions. By combining systematic field and laboratory analyses based on multiple analytical and statistical approaches, we explore the determinants of priming intensity along a 2200 km grassland transect on the Tibetan Plateau. Our results show that SOM stability characterized by chemical recalcitrance and physico-chemical protection explains more variance in the priming effect than plant, soil and microbial properties. High priming intensity (up to 137% of basal respiration) is associated with complex SOM chemical structures and low mineral-organic associations. The dependence of priming effect on SOM stabilization mechanisms should be considered in Earth System Models to accurately predict soil C dynamics under changing environments.