Carolyn Mitchell

Downstairs drivers - root herbivores shape communities of above-ground herbivores and natural enemies via changes in plant nutrients

1. Terrestrial food webs are woven from complex interactions, often underpinned by plant-mediated interactions between herbivores and higher trophic groups. Below- and above-ground herbivores can influence one another via induced changes to a shared host plant, potentially shaping the wider community. However, empirical evidence linking laboratory observations to natural field populations has so far been elusive. 2. This study investigated how root-feeding weevils (Otiorhynchus sulcatus) influence different feeding guilds of herbivore (phloem-feeding aphids, Cryptomyzus galeopsidis, and leaf-chewing sawflies, Nematus olfaciens) in both controlled and field conditions. 3. We hypothesized that root herbivore-induced changes in plant nutrients (C, N, P and amino acids) and defensive compounds (phenolics) would underpin the interactions between root and foliar herbivores, and ultimately populations of natural enemies of the foliar herbivores in the field. 4. Weevils increased field populations of aphids by ca. 700%, which was followed by an increase in the abundance of aphid natural enemies. Weevils increased the proportion of foliar essential amino acids, and this change was positively correlated with aphid abundance, which increased by 90% on plants with weevils in controlled experiments. 5. In contrast, sawfly populations were 77% smaller during mid-June and adult emergence delayed by >14 days on plants with weevils. In controlled experiments, weevils impaired sawfly growth by 18%, which correlated with 35% reductions in leaf phosphorus caused by root herbivory, a previously unreported mechanism for above-ground-below-ground herbivore interactions. 6. This represents a clear demonstration of root herbivores affecting foliar herbivore community composition and natural enemy abundance in the field via two distinct plant-mediated nutritional mechanisms. Aphid populations, in particular, were initially driven by bottom-up effects (i.e. plant-mediated effects of root herbivory), but consequent increases in natural enemies triggered top-down regulation.

Raspberry leaf blotch virus, a putative new member of the genus Emaravirus, encodes a novel genomic RNA

A new, segmented, negative-strand RNA virus with morphological and sequence similarities to other viruses in the genus Emaravirus was discovered in raspberry plants exhibiting symptoms of leaf blotch disorder, a disease previously attributed to the eriophyid raspberry leaf and bud mite (Phyllocoptes gracilis). The virus, tentatively named raspberry leaf blotch virus (RLBV), has five RNAs that each potentially encode a single protein on the complementary strand. RNAs 1, 2 and 3 encode, respectively, a putative RNA-dependent RNA polymerase, a glycoprotein precursor and the nucleocapsid. RNA4 encodes a protein with sequence similarity to proteins of unknown function that are encoded by the genomes of other emaraviruses. When expressed transiently in plants fused to green or red fluorescent protein, the RLBV P4 protein localized to the peripheral cell membrane and to punctate spots in the cell wall. These spots co-localized with GFP-tagged tobacco mosaic virus 30K cell-to-cell movement protein, which is itself known to associate with plasmodesmata. These results suggest that the P4 protein may be a movement protein for RLBV. The fifth RLBV RNA, encoding the P5 protein, is unique among the sequenced emaraviruses. The amino acid sequence of the P5 protein does not suggest any potential function; however, when expressed as a GFP fusion, it localized as small aggregates in the cytoplasm near to the periphery of the cell.

Plant Defense against Herbivorous Pests: Exploiting Resistance and Tolerance Traits for Sustainable Crop Protection

Interactions between plants and insect herbivores are important determinants of plant productivity in managed and natural vegetation. In response to attack, plants have evolved a range of defenses to reduce the threat of injury and loss of productivity. Crop losses from damage caused by arthropod pests can exceed 15% annually. Crop domestication and selection for improved yield and quality can alter the defensive capability of the crop, increasing reliance on artificial crop protection. Sustainable agriculture, however, depends on reduced chemical inputs. There is an urgent need, therefore, to identify plant defensive traits for crop improvement. Plant defense can be divided into resistance and tolerance strategies. Plant traits that confer herbivore resistance typically prevent or reduce herbivore damage through expression of traits that deter pests from settling, attaching to surfaces, feeding and reproducing, or that reduce palatability. Plant tolerance of herbivory involves expression of traits that limit the negative impact of herbivore damage on productivity and yield. Identifying the defensive traits expressed by plants to deter herbivores or limit herbivore damage, and understanding the underlying defense mechanisms, is crucial for crop scientists to exploit plant defensive traits in crop breeding. In this review, we assess the traits and mechanisms underpinning herbivore resistance and tolerance, and conclude that physical defense traits, plant vigor and herbivore-induced plant volatiles show considerable utility in pest control, along with mixed species crops. We highlight emerging approaches for accelerating the identification of plant defensive traits and facilitating their deployment to improve the future sustainability of crop protection.

Biology of the European large raspberry aphid (Amphorophora idaei): its role in virus transmission and resistance breakdown in red raspberry

1 The European large raspberry aphid Amphorophora idaei Börner is the most important vector of viral diseases afflicting commercially grown red raspberry (Rubus idaeus L.) in Northern Europe, with European raspberry production amounting to 416 000 tonnes per annum. This review synthesizes existing knowledge on its biology and interactions with other organisms, including its host plant and the viral pathogens it vectors.

Arthropod pests of currant and gooseberry crops in the U.K.: their biology, management and future prospects

1 Approximately 10–12 species of Ribes plants are cultivated for fruit production, mainly blackcurrants, red‐ and whitecurrants and gooseberries. These crops are increasingly recognized as rich sources of vitamin C and anthocyanins, with production rising by 24% in Europe subsequent to 1998. To date, research into insect pests of Ribes has been fragmented, with little appreciation of how changes in climate and agronomic practices affect biology. 2 We review 12 key pests of currant and gooseberry crops in Northern Europe, with specific emphasis on their biology and current management options. These are blackcurrant leaf curling midge Dasineura tetensi, blackcurrant sawfly Nematus olfaciens, common gooseberry sawfly Nematus ribesii, European permanent currant aphid Aphis schneideri, redcurrant blister aphid Cryptomyzus ribis, currant–sowthistle aphid Hyperomyzus lactucae, European gooseberry aphid Aphis grossulariae, woolly vine scale Pulvinaria vitis, common green capsid Lygocoris pabulinus, winter moth Operophtera brumata, clear wing moth Synanthedon tipuliformis and blackcurrant gall mite Cecidophyopsis ribis. 3 It is anticipated that global climate change could lead to increases in the incidence of some aphids through increased overwintering survival and longer seasonal activity. Moreover, changes in management practices such as increased cropping densities (from 5400 ha−1 to 8700 ha−1) and machine harvesting could lead to pest outbreaks through optimal microhabitats and increased susceptibility to pest colonization. 4 Future management options are considered, focusing on integrated pest management approaches, including behaviour‐manipulating semiochemicals, predictive models, biocontrol and improved plant resistance through breeding.

Combining plant resistance and a natural enemy to control Amphorophora idaei

The European large raspberry aphid Amphorophora idaei Börner (Homoptera: Aphididae) is a virus vector of at least four plant virus complexes making it the most important aphid pest of raspberries in Northern Europe. An approach combining a bottom-up control (plant resistance) and a top-down control (an aphid parasitoid) using Aphidius ervi Haliday (Hymenoptera: Aphidiinae) was investigated in the laboratory. Aphid performance (pre-reproductive period, total reproductive output, lifespan and rm) were compared when reared on both a susceptible cultivar and a resistant cultivar with significantly poorer performance on the resistant cultivar. Parasitoid attack behaviour increased with aphid density on both cultivars, but was significantly lower on resistant plants than susceptible plants. Aphids showed a greater tendency to drop from the plant when feeding on resistant plants compared with susceptible plants. The significance of the results is discussed in the context of possible control of the aphid using these combined methods.

Protected raspberry production accelerates onset of oviposition by vine weevils (Otiorhynchus sulcatus)

1 Soft fruit production is increasingly reliant on crops that are grown under the protection of plastic tunnels, which may also affect insect communities as a result of localized climate change and changes to host plant physiology and chemistry. In particular, insect development rates may differ from field populations, making it more difficult to target control measures. 2 The present study investigated how protected environments affected adult vine weevil (Otiorhynchus sulcatus) feeding and reproduction on red raspberry (Rubus idaeus). We focused on the period between adult emergence and the onset of oviposition (i.e. the pre‐reproductive period), which represents the optimal period for control. 3 Tunnels were up to 4 °C warmer than field plantations in 2008, with plants growing significantly faster (50% increase in height and 16% increase in leaf area) than field grown plants. The carbon/nitrogen ratio in leaves was higher in tunnels (12.07) than the field (10.89) as a result of a significant decrease in nitrogen concentrations (3.40 and 3.90 mg g−1, respectively). 4 Over 4 weeks, weevils consumed significantly more foliage in tunnels (370.89 mg) than weevils in the field (166.68 mg), suggesting compensatory feeding to counteract lower leaf nitrogen concentrations. Weevils in tunnels achieved sexual maturity 8 days earlier than those in the field and produced 20‐fold more eggs by the time they were 5 weeks old. 5 Applying a degree‐day model showed good agreement between predicted and observed pre‐reproductive periods for weevils in tunnels (36 and 30 days, respectively) and in field plots (41 and 38 days, respectively).

Protection of Pea Aphids Associated with Coinfecting Bacterial Symbionts Persists During Superparasitism by a Braconid Wasp

Bacterial endosymbionts that associate facultatively with insect herbivores can influence insect fitness and trophic interactions. The pea aphid, Acyrthosiphon pisum, can be protected from parasitism by the braconid wasp Aphidius ervi when harbouring particular symbiotic bacteria, with specific endosymbiont coinfections providing almost complete protection. However, studies often quantify aphid mummification with no control over parasitoid oviposition per aphid; thus, if mummy production fails or is low, the causes are often unclear. Here, we show that the high level of protection associated with the coinfecting endosymbionts Hamiltonella defensa and X-type is maintained even when pea aphids are superparasitised. This contrasts strongly with the protection provided by H. defensa alone, which has been shown by others to be overcome by superparasitism. By dissecting aphids exposed to two parasitoid attacks, we reveal that A. ervi deposits eggs equally freely in endosymbiont-infected and uninfected nymphs, and lack of mummification in endosymbiont-protected nymphs arises from failure of the wasp eggs to hatch or emerging larvae to develop.