Responses of pea plants to multiple antagonists are mediated by order of attack and phytohormone crosstalk.

Plants are often attacked by multiple antagonists and traits of the attacking organisms and their order of arrival onto hosts may affect plant defences. However, few studies have assessed how multiple antagonists, and varying attack order, affect plant defence or nutrition. To address this, we assessed defensive and nutritional responses of Pisum sativum plants after attack by a vector herbivore (Acrythosiphon pisum), a nonvector herbivore (Sitona lineatus), and a pathogen (Pea enation mosaic virus, PEMV). We show viruliferous A. pisum induced several antipathogen plant defence signals, but these defences were inhibited by S. lineatus feeding on peas infected with PEMV. In contrast, S. lineatus feeding induced antiherbivore defence signals, and these plant defences were enhanced by PEMV. Sitona lineatus also increased abundance of plant amino acids, but only when they attacked after viruliferous A. pisum. Our results suggest that diverse communities of biotic antagonists alter defence and nutritional traits of plants through complex pathways that depend on the identity of attackers and their order of arrival onto hosts. Moreover, we show interactions among a group of biotic stressors can vary along a spectrum from antagonism to enhancement/synergism based on the identity and order of attackers, and these interactions are mediated by a multitude of phytohormone pathways.

Vector-borne plant pathogens modify top-down and bottom-up effects on insect herbivores.

Ecological theory predicts that host-plant traits affect herbivore population growth rates, which in turn modulates predator-prey interactions. However, while vector-borne plant pathogens often alter traits of both host plants and vectors, a few studies have assessed how pathogens may act as interaction modifiers within tri-trophic food webs. By applying a food web motif framework, we assessed how a vector-borne plant pathogen (Pea-enation mosaic virus, PEMV) modified both bottom-up (plant-herbivore) and top-down (predator-prey) interactions. Specifically, we assessed trophic interactions with PEMV-infectious Acyrthosiphon pisum (pea aphid) vectors compared to non-infectious aphids in a factorial experiment that manipulated predator and plant communities. We show that PEMV altered bi-trophic relationships, whereby on certain plant species, PEMV reduced vector performance but also increased their susceptibility to predators. However, on other plant species, PEMV weakened top-down control or increased vector performance. Our results suggest that vector-borne plant pathogens are important interaction modifiers for plant-herbivore-predator dynamics: host-plant response to viruses can decrease herbivore abundance by reducing herbivore performance, but also increase herbivore abundance by weakening top-down control. Broadly speaking, trophic interactions that regulate herbivore outbreaks appear to be modified for herbivores actively transmitting viruses to host plants. Consequently, management and monitoring of outbreaking herbivores should consider the infection status of focal populations.

Plant-mediated interactions between a vector and a non-vector herbivore promote the spread of a plant virus.

Herbivores that transmit plant pathogens often share hosts with non-vector herbivores. These co-occurring herbivores can affect vector fitness and behaviour through competition and by altering host plant quality. However, few studies have examined how such interactions may both directly and indirectly influence the spread of a plant pathogen. Here, we conducted field and greenhouse trials to assess whether a defoliating herbivore (Sitona lineatus) mediated the spread of a plant pathogen, Pea enation mosaic virus (PEMV), by affecting the fitness and behaviour of Acrythosiphon pisum, the PEMV vector. We observed higher rates of PEMV spread when infectious A. pisum individuals shared hosts with S. lineatus individuals. Using structural equation models, we showed that herbivory from S. lineatus increased A. pisum fitness, which stimulated vector movement and PEMV spread. Moreover, plant susceptibility to PEMV was indirectly enhanced by S. lineatus, which displaced A. pisum individuals to the most susceptible parts of the plant. Subsequent analyses of plant defence genes revealed considerable differences in plant phytohormones associated with anti-herbivore and anti-pathogen defence when S. lineatus was present. Given that vectors interact with non-vector herbivores in natural and managed ecosystems, characterizing how such interactions affect pathogens would greatly enhance our understanding of disease ecology.

The beneficial effect of prior experience on the acquisition of spatial memory in rats with CA1, but not large hippocampal lesions: a possible role for schema formation.

Prior experience has been shown to improve learning in both humans and animals, but it is unclear what aspects of recent experience are necessary to produce beneficial effects. Here, we examined the capacity of rats with complete hippocampal lesions, restricted CA1 lesions, or sham surgeries to benefit from prior experience. Animals were tested in two different spatial tasks in the watermaze, the conventional watermaze task and delayed match-to-position. The two lesions impaired performance in both tasks when rats had no prior experience. However, when given prior training with one task, CA1 lesions had no effect on performance in the other task. In contrast, rats with hippocampal lesions did not benefit from prior training. The findings show that prior experience can benefit learning even when the previously learned task and a new task are quite different. The concept of schema may be useful for understanding the benefits of prior experience.

Recent and remote retrograde memory deficit in rats with medial entorhinal cortex lesions.

The hippocampus is critically involved in the acquisition and retrieval of spatial memories. Even though some memories become independent of the hippocampus over time, expression of spatial memories have consistently been found to permanently depend on the hippocampus. Recent studies have focused on the adjacent medial entorhinal cortex (MEC), as it provides major projections to the hippocampus. These studies have shown that lesions of the MEC disrupt spatial processing in the hippocampus and impair spatial memory acquisition on the watermaze task. MEC lesions acquired after learning the watermaze task also disrupt recently acquired spatial memories. However, the effect of MEC lesions on remotely acquired memories is unknown. The current study examined the effect of MEC lesions on recent and remote memory retrieval using three hippocampus-dependent tasks: the watermaze, trace fear conditioning, and novel object recognition. MEC lesions caused impaired retrieval of recently and remotely acquired memory for the watermaze. Rats with MEC lesions also showed impaired fear memory when exposed to the previously conditioned context or the associated tone, and this reduction was seen both when the lesion occurred soon after trace fear condition and when it occurred a month after conditioning. In contrast, MEC lesions did not disrupt novel object recognition. These findings indicate that even with an intact hippocampus, rats with MEC lesions cannot retrieve recent or remote spatial memories. In addition, the involvement of the MEC in memory extends beyond is role in navigation and place memory.

Keystone mutualism strengthens top-down effects by recruiting large-bodied ants.

Determining the impacts of mutualistic interactions and predator diversity on food webs are two important goals in community ecology. In this study, we examined how predator community variation mediates the strength of top-down effects in the presence and absence of mutualistic interactions. We examined the impacts of predatory ant species that simultaneously prey on leaf-chewing herbivores (Lepidoptera) and engage in food-for-protection mutualisms with sap-feeding herbivores (Hemiptera) in the lower canopy of Connecticut forests. In this 2-year study, we examined three hypothetical mechanisms by which mutualisms can alter the top-down effects of ants: (1) sap feeders increase ant abundance, thus strengthening predatory effects; (2) sap feeders increase the relative abundance of a species that has stronger predatory effects; and (3) changes to predator diversity (species richness) are caused by sap feeders mediating top-down effects of the ant community. Experiments revealed that host plants occupied by sap feeders favored large-bodied ant species in the genus Camponotus, but there were no changes to community-wide ant abundance or ant species richness. Fitting predictions of predation strength based on the functional trait of body size, large-bodied Camponotus suppressed caterpillars and reduced leaf herbivory. This work shows that the ant-hemipteran mutualism, which has been characterized as a keystone interaction, can generate strong top-down effects on leaf-chewing herbivores and herbivory via increasing the relative abundance of species with functional traits relevant to predation, such as body size. Therefore, the emergence of specific ants as keystone predators in a community can be contingent upon their mutualism with sap-feeding Hemiptera.

Hippocampal area CA1 and remote memory in rats.

Hippocampal lesions often produce temporally graded retrograde amnesia (TGRA), whereby recent memory is impaired more than remote memory. This finding has provided support for the process of systems consolidation. However, temporally graded memory impairment has not been observed with the watermaze task, and the findings have been inconsistent with context fear conditioning. One possibility is that large hippocampal lesions indirectly disrupt (by retrograde degeneration) the function of areas that project to the hippocampus that are important for task performance or thought to be important for storing consolidated memories. We developed a discrete lesion targeting area CA1, the sole output of the hippocampus to neocortex, and tested the effects of this lesion on recent and remote memory in the watermaze task, in context fear conditioning, and in trace fear conditioning. In all three tasks, recent and remote memory were similarly impaired after CA1 lesions. We discuss factors that help to illuminate these findings and consider their relevance to systems consolidation.

Predatory birds and ants partition caterpillar prey by body size and diet breadth.

The effects of predator assemblages on herbivores are predicted to depend critically on predator-predator interactions and the extent to which predators partition prey resources. The role of prey heterogeneity in generating such multiple predator effects has received limited attention. Vertebrate and arthropod insectivores constitute two co-dominant predatory taxa in many ecosystems, and the emergent properties of their joint effects on insect herbivores inform theory on multiple predator effects as well as biological control of insect herbivores. Here we use a large-scale factorial manipulation to assess the extent to which birds and ants engage in antagonistic predator-predator interactions and the consequences of heterogeneity in herbivore body size and diet breadth (i.e. the diversity of host plants used) for prey partitioning. We excluded birds and reduced ant density (by 60%) in the canopies of eight northeastern USA deciduous tree species during two consecutive years and measured the community composition and traits of lepidopteran larvae (caterpillars). Birds did not affect ant density, implying limited intraguild predation between these taxa in this system. Birds preyed selectively upon large-bodied caterpillars (reducing mean caterpillar length by 12%) and ants preyed selectively upon small-bodied caterpillars (increasing mean caterpillar length by 6%). Birds and ants also partitioned caterpillar prey by diet breadth. Birds reduced the frequency dietary generalist caterpillars by 24%, while ants had no effect. In contrast, ants reduced the frequency of dietary specialists by 20%, while birds had no effect, but these effects were non-additive; under bird exclusion, ants had no detectable effect, while in the presence of birds, they reduced the frequency of specialists by 40%. As a likely result of prey partitioning by body size and diet breadth, the combined effects of birds and ants on total caterpillar density were additive, with birds and ants reducing caterpillar density by 44% and 20% respectively. These results show evidence for the role of prey heterogeneity in driving functional complementarity among predators and enhanced top-down control. Heterogeneity in herbivore body size and diet breadth, as well as other prey traits, may represent key predictors of the strength of top-down control from predator communities.

Distinct roles of hippocampus and medial prefrontal cortex in spatial and nonspatial memory.

In earlier work, patients with hippocampal damage successfully path integrated, apparently by maintaining spatial information in working memory. In contrast, rats with hippocampal damage were unable to path integrate, even when the paths were simple and working memory might have been expected to support performance. We considered possible ways to understand these findings. We tested rats with either hippocampal lesions or lesions of medial prefrontal cortex (mPFC) on three tasks of spatial or nonspatial memory: path integration, spatial alternation, and a nonspatial alternation task. Rats with mPFC lesions were impaired on both spatial and nonspatial alternation but performed normally on path integration. By contrast, rats with hippocampal lesions were impaired on path integration and spatial alternation but performed normally on nonspatial alternation. We propose that rodent neocortex is limited in its ability to construct a coherent spatial working memory of complex environments. Accordingly, in tasks such as path integration and spatial alternation, working memory cannot depend on neocortex alone. Rats may accomplish many spatial memory tasks by relying on long-term memory. Alternatively, they may accomplish these tasks within working memory through sustained coordination between hippocampus and other cortical brain regions such as mPFC, in the case of spatial alternation, or parietal cortex in the case of path integration. © 2016 Wiley Periodicals, Inc.

Multiple interaction types determine the impact of ant predation of caterpillars in a forest community.

Direct and indirect effects of predators are highly variable in complex communities, and understanding the sources of this variation is a research priority in community ecology. Recent evidence indicates that herbivore community structure is a primary determinant of predation strength and its cascading impacts on plants. In this study, we use variation in herbivore community structure among plant species to experimentally test two hypotheses in a temperate forest food web. First, variation in the strength of predator effects, such as ant predation of caterpillars, is predicted to be density dependent, exhibiting stronger effects when prey abundance is high (density-dependent predation hypothesis). Second, mutualistic interactions between ants and sap-feeding herbivores are expected to increase the abundance of predatory ants, strengthening predation effects on herbivores with cascading effects on host plants (keystone mutualism hypothesis). Using a large-scale predator exclusion experiment across eight dominant tree species, we tracked changes in insect density on 862 plants across two years, recording 2,322 ants, 1,062 sap-feeders, 5,322 caterpillars, and quantifying herbivory on 199, 338 leaves. In this experiment, density-dependent predation did not explain variation in the direct or indirect effects of ants on caterpillars and herbivory. In partial support of the keystone mutualism hypothesis, sap-feeders strengthened top-down effects of ants on caterpillars under some conditions. However, stronger ant predation of caterpillars did not lead to measurable trophic cascades on trees occupied by sap-feeders. Instead, the presence of sap-feeders was associated with increased per capita feeding damage by caterpillars, and this bottom-up effect attenuated the indirect effects of ants on host plants. These findings demonstrate that examining the multi-trophic impacts of mutualisms and predation in the context of the broader community can reveal patterns otherwise masked by compensatory interactions.

Consolidation of spatial memory in the rat: Findings using zeta-inhibitory peptide.

Whether or not spatial memories reorganize in the rodent brain is an unanswered question that carries the importance of whether the rodent provides a suitable animal model of human retrograde amnesia. The finding of equally impaired recent and remote spatial memory could reflect the continued importance of the hippocampus for spatial memory or a performance deficit (for example, hippocampal lesions may impair the rat's ability to use distal spatial cues to navigate to a specific point in space). In the current study, we tested recent and remote spatial memory in rats following hippocampal ZIP (zeta-pseudosubstrate inhibitory peptide) infusion to inhibit PKMzeta. Hippocampal ZIP infusion has previously been shown to impair spatial and nonspatial memory soon after learning, presumably by reversing late-phase long-term potentiation, allowing us to disrupt memory without damaging hippocampal tissue. We used a stereotaxic approach for infusing ZIP throughout the dorsal, intermediate, and ventral hippocampus following spatial memory training. Although rats showed intact memory retrieval on the standard Morris watermaze task and trace fear conditioning, rats infused with ZIP 24h after training on the annular watermaze task exhibited impaired spatial memory compared to control rats (those infused with aCSF) and performed no different than chance. In contrast, rats infused with ZIP 1month after training performed similar to control rats and both groups performed above chance. Additionally, the ability to form new memories after ZIP infusions remained intact. Thus, ZIP infusions into the hippocampus after learning impaired retrieval of recently formed spatial memories while sparing remote spatial memories.

Transition from Beam-Target to Thermonuclear Fusion in High-Current Deuterium Z-Pinch Simulations.

Fusion yields from dense, Z-pinch plasmas are known to scale with the drive current, which is favorable for many potential applications. Decades of experimental studies, however, show an unexplained drop in yield for currents above a few mega-ampere (MA). In this work, simulations of DD Z-Pinch plasmas have been performed in 1D and 2D for a constant pinch time and initial radius using the code Lsp, and observations of a shift in scaling are presented. The results show that yields below 3 MA are enhanced relative to pure thermonuclear scaling by beamlike particles accelerated in the Rayleigh-Taylor induced electric fields, while yields above 3 MA are reduced because of energy lost by the instability and the inability of the beamlike ions to enter the pinch region.

Similarity in form and function of the hippocampus in rodents, monkeys, and humans.

We begin by describing an historical scientific debate in which the fundamental idea that species are related by evolutionary descent was challenged. The challenge was based on supposed neuroanatomical differences between humans and other primates with respect to a structure known then as the hippocampus minor. The debate took place in the early 1860 s, just after the publication of Darwin's famous book. We then recount the difficult road that was traveled to develop an animal model of human memory impairment, a matter that also turned on questions about similarities and differences between humans and other primates. We then describe how the insight that there are multiple memory systems helped to secure the animal model and how the animal model was ultimately used to identify the neuroanatomy of long-term declarative memory (sometimes termed explicit memory). Finally, we describe a challenge to the animal model and to cross-species comparisons by considering the case of the concurrent discrimination task, drawing on findings from humans and monkeys. We suggest that analysis of such cases, based on the understanding that there are multiple memory systems with different properties, has served to emphasize the similarities in memory function across mammalian species.

Contrasting effects on path integration after hippocampal damage in humans and rats.

The hippocampus and other medial temporal lobe structures have been linked to both memory and spatial cognition, but it has been unclear how these ideas are connected. We carried out parallel studies of path integration in patients with medial temporal lobe lesions and rats with hippocampal lesions. Subjects entered a circular arena without vision, searched for a target, and then attempted to return to the start location. Patients performed accurately, and as well as controls, so long as the outward path was relatively direct and the target was found within 20 s. In sharp contrast, rats with hippocampal lesions were impaired, even when the outward path was shorter than 1 m, involved no turns, and the target was found within 3 s. We suggest that patients succeeded because performance could be supported by working memory and that patients and rats differ after hippocampal lesions in their ability to construct a coherent working memory of spatial environments.

Hippocampus, perirhinal cortex, and complex visual discriminations in rats and humans.

Structures in the medial temporal lobe, including the hippocampus and perirhinal cortex, are known to be essential for the formation of long-term memory. Recent animal and human studies have investigated whether perirhinal cortex might also be important for visual perception. In our study, using a simultaneous oddity discrimination task, rats with perirhinal lesions were impaired and did not exhibit the normal preference for exploring the odd object. Notably, rats with hippocampal lesions exhibited the same impairment. Thus, the deficit is unlikely to illuminate functions attributed specifically to perirhinal cortex. Both lesion groups were able to acquire visual discriminations involving the same objects used in the oddity task. Patients with hippocampal damage or larger medial temporal lobe lesions were intact in a similar oddity task that allowed participants to explore objects quickly using eye movements. We suggest that humans were able to rely on an intact working memory capacity to perform this task, whereas rats (who moved slowly among the objects) needed to rely on long-term memory.

Hippocampal Infusion of Zeta Inhibitory Peptide Impairs Recent, but Not Remote, Recognition Memory in Rats.

Spatial memory in rodents can be erased following the infusion of zeta inhibitory peptide (ZIP) into the dorsal hippocampus via indwelling guide cannulas. It is believed that ZIP impairs spatial memory by reversing established late-phase long-term potentiation (LTP). However, it is unclear whether other forms of hippocampus-dependent memory, such as recognition memory, are also supported by hippocampal LTP. In the current study, we tested recognition memory in rats following hippocampal ZIP infusion. In order to combat the limited targeting of infusions via cannula, we implemented a stereotaxic approach for infusing ZIP throughout the dorsal, intermediate, and ventral hippocampus. Rats infused with ZIP 3-7 days after training on the novel object recognition task exhibited impaired object recognition memory compared to control rats (those infused with aCSF). In contrast, rats infused with ZIP 1 month after training performed similar to control rats. The ability to form new memories after ZIP infusions remained intact. We suggest that enhanced recognition memory for recent events is supported by hippocampal LTP, which can be reversed by hippocampal ZIP infusion.

Tracking flight activity of potato leafhopper (Hemiptera: Cicadellidae) with the Midwest Suction Trap Network.

Potato leafhopper (PLH), Empoasca fabae Harris (Hemiptera: Cicadellidae), is an economic pest of a variety of crops that migrates between overwintering sites in the southern United States and northern breeding grounds. Since 2005, the Midwest Suction Trap Network (STN) has monitored the magnitude and timing of aerially dispersing aphids' activity, but the potential of the network to monitor other taxa is only beginning to be explored. Here, we use the Midwest STN to examine how the magnitude and timing of PLH activity vary with weather, cropland cover, and time of year. We found that weekly PLH activity increased early in the season (May-June) with increasing degree day accumulation and decreased mid-season (July-August) with increasing occurrence of rain. The first detections occurred earlier in southern latitudes, while the last detections occurred sooner, when there was more surrounding potato land cover, and later over time between 2018 and 2021 and in southern latitudes. PLH activity was thus longer in duration in southern latitudes and has continued to extend later into the year overall. Resolving uncertainty about how well the Midwest STN captures migratory activity and how closely suction trap detections reflect local population densities in crop fields remain important research priorities before the potential of the Midwest STN for PLH monitoring can be realized. Still, observed patterns suggest that PLH could increase in economic importance as insects disperse over larger portions of the growing season in the warming, agriculturally productive US Midwest and that the STN can become a useful tool to monitor these changes.

Remote context fear conditioning remains hippocampus-dependent irrespective of training protocol, training-surgery interval, lesion size, and lesion method.

Systems consolidation involves the reorganization of brain circuits that support long-term memory. It is a prolonged process that can take days, weeks, or longer. An animal model of systems consolidation was established in the early 1990s and provided compelling support for the initial observations in humans, that hippocampal damage disproportionally impairs recent memory compared to remote memory. Context fear conditioning was the most frequently and successfully used task to study systems consolidation and demonstrate temporally graded retrograde amnesia. However, recent studies have failed to support these early findings of temporal gradients and instead reported that both recent and remote memories are equally impaired. Thus, the status of context fear conditioning as method to study the process of systems consolidation is at present uncertain. Accordingly, we evaluated classically conditioned fear memory in large groups of rats with hippocampal damage by manipulating several procedural variables including the training protocol, the training-surgery interval, the extent of hippocampal damage, and the method of damaging the hippocampus. The results indicate that hippocampal damage profoundly impairs context fear conditioning. These findings are unambiguous and independent of any particular procedural manipulation we evaluated. We suggest that the preponderance of currently available evidence indicates that context fear memory remains hippocampus-dependent indefinitely.

Temporal and spatial dynamics of the emerald ash borer invasion in Connecticut as shown by the native digging wasp Cerceris fumipennis (Hymenoptera: Crabronidae).

Detecting and monitoring populations of the invasive emerald ash borer (EAB) is crucial to successful management of the pest and evaluation of its ecological impacts. However, the beetle's cryptic habit makes accurate monitoring costly and time-consuming. Biosurveillance takes advantage of the foraging effort of a predatory wasp Cerceris fumipennis (Hymenoptera: Crabronidae). This native, solitary, ground-nesting hunting wasp hunts adult buprestid beetles to provision its brood cells. By intercepting the hunting wasps, we can learn which species of buprestids are in the surrounding forest. The resulting data provides information on the presence and relative abundance of invasive buprestids like EAB which can supplement other monitoring efforts. In this paper we share results of ten years of biosurveillance surveys of the EAB in Connecticut. Among 112 sites, we observed EAB populations; from first detection, through the population peak and then through to the population crash, matching patterns observed in other regions of the United States. We also observed the spread of the EAB relative abundance as it moved through the state following an invasion front starting in New Haven, Co. The average time from first detection to population crash was nine years. On average, populations peaked three years after first detection, and remained at peak levels for three to four years. Population decline was gradual and took another three to four years. Notably, no evidence of a second introduction to Connecticut was seen with proportional abundance increasing over time after expanding outward from the introduction point. These results corroborate other traditional monitoring efforts in the eastern U.S. and provide independent validation of predicted population dynamics in ash stands.