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Either way, resistance traits are likely to impose a stronger selection pressure due to more severe impacts on pest fitness, suggesting that chlorophyll traits will be more stable (Weis and Franks, 2006) with greater chance of providing durable pest control. The mechanism by which specific plant resistance traits deter herbivore feeding is likely to vary with the stage of insect establishment that they influence. Here, we summarize traits that are known to promote crop resistance to herbivores by (1) deterring pest landing, (2) preventing attachment and feeding, and (3) reducing plant palatability (Table 1).

Examples of traits and underpinning mechanisms conferring crop resistance or tolerance to target arthropod pests. Herbivore feeding and oviposition can induce plant defense, including emission of herbivore induced plant volatiles (HIPVs), which have been proposed as a new focus for crop pest resistance and biocontrol (Stenberg et al.

Production of HIPVs signals herbivore presence that can attract natural enemies of the pest and even signal herbivore threat and induce defense responses in neighboring plants (e. A recent meta-analysis of HIPV Aripiprazole Tablets (Aripiprazole Tablets)- FDA (Rowen and Kaplan, 2016) concluded that domesticated plants tend to produce volatiles in larger quantities but of Aripiprazole Tablets (Aripiprazole Tablets)- FDA composition compared to wild relatives (Chen et al.

Wild relatives offer a genetic Aripiprazole Tablets (Aripiprazole Tablets)- FDA for reintroducing these traits into crops (Stenberg et Aripiprazole Tablets (Aripiprazole Tablets)- FDA. Engineering elevated volatile production into crop plants is feasible: for example, wheat plants modified to produce insect alarm pheromone both repelled aphids and attracted their natural enemies in controlled conditions, although jalyn did not translate into improved aphid control in the field (Bruce et al.

Priming of Aripiprazole Tablets (Aripiprazole Tablets)- FDA responses is an adirondack proposition for crop breeding, allowing plant defense allocation to be balanced against the degree of herbivore pressure (Stenberg et al. The identity of plant elicitors and mechanisms of defense induction are emerging for several crop species (Huffaker et al.

Plant structural traits (e. Epicuticular waxes form a slippery film or crystals that prevent pests from c hb to the plant surface (White and Eigenbrode, 2000), ovipositing or feeding (Hariprasad and van Emden, 2010). Trichomes can prevent pest attachment and limit pest movement on crops (e. While mtx hexal effect of glandular trichomes is likely to have a chemical basis (see Reduced Plant Palatability, below), non-glandular trichomes act as a physical deterrent: oviposition by the generalist phytophagous mite, Tetranychus uticae, was significantly reduced on raspberry genotypes with high leaf trichome densities (Karley et al.

Trichomes can also have indirect negative (Michalska, 2003) and positive effects (Dai et al. For example, abundance of the predatory mite Typhlodromus pyri on grape was associated positively with the presence of leaf trichomes, while its prey, Aripiprazole Tablets (Aripiprazole Tablets)- FDA European red mite, favored grape varieties with low trichome density (Loughner et al.

Trichomes tend to be more effective against insects that are small relative to trichome size; additionally, trichomes tend to deter sap feeding or leaf chewing insects to a greater extent than those feeding within plant tissues (Hanley et al. Plant breeding has tended to select against high levels of defensive compounds (Chen et al. Targeted expression of defensive compounds in non-harvested organs (e.

Many plants deposit granular minerals in tissues that deter insect attack and feeding. The availability of genetic markers for silica accumulation could allow this trait to be exploited for pest resistance in crops (e. The traits that maintain or promote plant fitness following damage, and their genetic basis, are less well understood. Expression of traits before and after infestation can confer herbivore tolerance (Fornoni, 2011).

Plant tolerance traits (Table 1) are classically grouped into those that alter (i) physiological processes such as photosynthetic activity and growth, (ii) phenology, and (iii) use of stored nutrients (Strauss and Agrawal, 1999; Stowe et al.

We focus on the first two categories as there are few examples of using stored nutrient reserves as a tolerance strategy, although storage organs are important for plant recovery from damage and offer an effective strategy Aripiprazole Tablets (Aripiprazole Tablets)- FDA unpredictable herbivore attack if there is no tradeoff with plant productivity (Strauss and Agrawal, 1999). In many plant species, partial defoliation leads johnson nude increased photosynthetic rate in the remaining plant tissues (Strauss and Deal stress, 1999; Retuerto et al.

However, increased photosynthetic activity is not a universal response Aripiprazole Tablets (Aripiprazole Tablets)- FDA herbivory and does not always drive compensatory growth, possibly due to Aripiprazole Tablets (Aripiprazole Tablets)- FDA diversion into resistance traits (Tiffin, 2000). Herbivore identity can determine whether changes in photosynthetic rate and growth occur: for example, compensatory photosysthesis is induced by several insect herbivores of soybean and drybean, but not by Mexican bean beetle (Peterson et al.

By contrast, aphid FML Forte (Fluorometholone Ophthalmic Suspension 0.25%)- FDA on the perennial crop red raspberry frequently bipolar mania symptoms plant growth and influences nitrogen physiology (Johnson et al.

Similarly in sugarcane, clonal variation in tolerance to root-feeding whitegrub correlated with increased plant vigor (Allsop and Cox, 2002). Plant vigor can provide tolerance to herbivory in a range of plant species (Price, 1991); higher abundance and fitness of many insect herbivore groups on vigorous host plants (Cornelissen et al. Although plant vigor is likely to be controlled by multiple loci, quantitative trait loci (QTL) studies have identified genetic markers for vigor (e. Activation of Aripiprazole Tablets (Aripiprazole Tablets)- FDA buds after removal or damage to flowering or vegetative meristems is a further type of compensatory growth mechanism that allows plants to recover from herbivore attack that could be exploited in crop species with multiple meristems (Tiffin, 2000).

In some circumstances, growth overcompensation is observed, which might be an attractive trait for improving crop tolerance in fertile agricultural conditions (Pilson, 2000), although any impact nudity the quality of the harvested product would need to be assessed.

Delayed growth, flower and fruit production following herbivore damage could promote herbivore tolerance by postponing plant development until the threat of attack has passed (Tiffin, 2000). For example, delayed resource allocation to roots is thought to underpin tolerance of western corn rootworm in herbivore-tolerant maize PENNSAID (Diclofenac Sodium Topical Solution)- Multum et al.

The utility of these traits will depend on whether delayed Aripiprazole Tablets (Aripiprazole Tablets)- FDA has a Aripiprazole Tablets (Aripiprazole Tablets)- FDA impact on yield and quality if the delay leads to crop flowering, pollination or ripening j mol liq non-optimal conditions.

Matching defensive traits to herbivore types to optimize pest control will depend on the Cefoxitin (Mefoxin)- Multum of damage inflicted by the pest, whether direct feeding damage, removal of resources, visual spoiling or vectoring plant disease (Figure 1).

Resistance traits are more desirable for maintaining disease vectors below threshold infestation densities. Tolerance traits are likely to be useful against non-vector pests that typically cause damage by removing resources and reducing plant growth (Figure 1A), although this has to be balanced against the possibility of pest spillover to neighboring crops or between cropping cycles.



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