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Water Deficit Stress - Host Plant Nutrient Accumulations and Associations with Phytophagous Arthropods

Written By

Allan T. Showler

Submitted: 01 May 2012 Published: 13 March 2013

DOI: 10.5772/53125

From the Edited Volume

Abiotic Stress - Plant Responses and Applications in Agriculture

Edited by Kourosh Vahdati and Charles Leslie

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1. Introduction

When the availability of water is insufficient to maintain plant growth, photosynthesis, and transpiration, plants become water deficit stressed (Fan et al., 2006), a serious problem that reduces world crop production (Boyer, 1982; Vincent et al., 2005). While drought has profound direct detrimental effects against plants, including rendering otherwise arable regions less, or non-, arable, herbivorous arthropod populations and the injuries they cause can be affected by stress-related changes that occur in the plant. Moderate stress is known to heighten the nutritional value of some plants’ tissues and juices, in some instances to reduce concentrations of plant defense compounds, and even to select against predators and parasitoids that otherwise help reduce pest populations to economically tolerable levels, each of which can contribute toward greater pest infestations. Sometimes the injury inflicted on water deficit stressed plants is intensified even if numbers of the pest haven’t been affected, as in the instances of honeylocust spider mites, Platytetranychus multidigituli (Ewing), on honeylocust trees, Gleditsia triacanthos L. (Smitley & Peterson, 1996), and greenbug and flea beetle, Aphtona euphorbiae Schrank, on several different crop species (Popov et al., 2006). When the stress associated with water deficit is more severe, however, host plant suitability for utilization by arthropods declines (Mattson & Haack, 1987; Showler, 2012 ) because of insufficient availability of water for the pest, and from senescence and drying of the plant’s tissues. As plants desiccate further, they eventually die and concerns about arthropod pest damage to that crop become moot unless the pests move from unsuitable dead plant material to vulnerable, living crops.

Although severe water deficit stress that causes plant mortality usually renders plants useless to herbivores, chronic lower level or pulsed water deficit stress can enhance the nutritional value of plants to arthropods, resulting in selection preference, heightened populations, intensified injury to crops, and even outbreaks that affect production on area-wide scales. Twospotted spider mite, Tetranychus urticae Koch, populations, for example, increase on drought stressed soybeans, Glycine max (L.) Merrill (Klubertanz et al., 1990) and populations of the Russian wheat aphid, Diuraphis noxia (Morvilko), increased in nonirrigated wheat, Triticum aestivum L., fields as compared with fields that received irrigation (Archer et al., 1995). The cabbage aphid, Brevicoryne brassicae L., infested water deficit stressed rape, Brassica napus L., more heavily than nonstressed plants (Burgess et al., 1994; Popov et al., 2006), and greenbug, Schizaphis graminum (Rondani), densities were higher and more injurious to wheat stressed by drought (Dorschner et al., 1986). Water deficit stressed host plants are also known to favor the xerophilic maize leaf weevil, Tanymecus dilaticollis Gyllenhall (Popov et al., 2006); scolytid bark beetles infesting trees (Lorio et al., 1995); flea beetles on corn, Zea mays L. (Bailey, 2000); and the fall armyworm, Spodoptera frugiperda (J. E. Smith), on tall fescue, Festuca arundinacea Schreb. (Bultman & Bell, 2003). Under circumstances where water deficit is beneficial to arthropod pests, population growth generally results in further damage to crops that have already been injured or stunted by water deficit stress itself.

Water deficit stress in plants can affect the amounts and composition of volatile compounds, and the concentrations of several kinds of nutrients beneficial to arthropod pests. Its associations with free amino acids and carbohydrates are chiefly described in this chapter because those two kinds of nutrients have been researched to an appreciable extent, permitting some conclusions to be drawn about arthropod host plant selection and levels of infestation.

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2. Water deficit, host plant nutrient accumulation, and associations with phytophagous arthropods

Water deficit stress alters plant metabolism and biochemistry (Hsiao, 1973; Beck et al., 2007), and consequent changes to plant physiological processes have been reported as being factors affecting herbivorous arthropod host plant preferences, growth, and development (Mattson & Haack, 1987; Showler, 2012 ). Although soil dries in association with drought, evapotranspiration rates in affected plants are often maintained (Jordan & Ritchie, 1971) by elevated accumulations of free amino acids, especially proline, and other organic solutes (Janagouar et al., 1983). Osmotic stress in plants involves several interlinked molecular pathways that transmit signals and produce stress-responsive metabolites (Ingram & Bartels, 1996; Zhu, 2002), and gene transcripts associated with signaling can be up- or down-regulated minutes after stress induction (Seki et al., 2001; Showler et al., 2007). Water deficit stressed plants often have diminished osmotic potential (Labanauskas et al., 1981; Golan-Goldhirsch et al., 1989; Bussis & Heineke, 1998), heightened oxidative stress (Becana et al., 1998; Knight & Knight, 2001), and accumulations of osmolytes such as antioxidants, amino acids, carbohydrates, and inorganic ions, altering the attractiveness and nutritional value of the plant (Jones, 1991; Showler & Castro, 2010a). Reduced leaf water content relative to dry biomass in water deficit stressed plants, in combination with the increased quantities of nutritional metabolites (White, 1984; Dubey, 1999; Ramanulu et al., 1999; Garg et al., 2001), may contribute toward the increased nutritional value of plants per unit of surface area consumed by arthropods. It is likely that arthropods can perceive cues about host plant suitability from emission of plant volatile compounds, or semiochemicals.

Chemical cues from plants play a major, perhaps decisive, role in host plant selection and utilization by herbivorous arthropods (Schur & Holdaway, 1970; Fenemore, 1980; Waladde, 1983; Burton & Schuster, 1981; Ramaswamy, 1988; Salama et al., 1984; Udayagiri & Mason, 1995). Water deficit stress in plants alters plant metabolism which can affect quantities and combinations of volatile compounds (Apelbaum & Yang, 1981; Hansen & Hitz, 1982; Zhang & Kirkham, 1990). Apple trees, Malus domestica Borkh., for instance, emit 29 volatile compounds, some of them in elevated amounts during water deficit stress (Ebel et al., 1995). Many phytophagous arthropods appear to respond to certain blends of volatiles (Miller & Strickler, 1984) that signal the host plant’s nutritional value (Mattson & Haack, 1987; Bernays & Chapman, 1994; Showler, 2012 ). Increased production of volatiles (e.g., ethylene, acetaldehyde, and ethanol) resulting from plant stress (Kimmerer & Kozlowski, 1982) can be attractive to some herbivorous arthropods and repellent to others (Chrominsky et al., 1982; Dunn et al., 1986; Haack & Slansky, 1987; Bernays & Chapman, 1994). Ethylene, for example, attracts the boll weevil, Anthonomus grandis grandis Boheman (Hedin et al., 1976), and, in many host plants it can increase susceptibility to the Egyptian cotton leafworm, Spodoptera littoralis Boisd. (Stotz et al., 2000), but ethylene deters the fall armyworm from corn (Harfouche et al., 2006) and the olive moth, Prays oleae Bern, from olive trees (Ramos et al., 2008). Forest outbreaks of many species of scolytid bark beetles (Hodger & Lorio, 1975; Wright et al., 1979; Vité et al., 1986; Ormeño et al., 2007; Branco et al., 2010) and the western spruce budworm, Choristoneura occidentalis Freeman, are related to amounts and kinds of host plant volatiles emitted during conditions of drought (Cates & Redak, 1988).

Once the phytophagous arthropod has found or selected the host plant, contact chemoreceptors on many are important in the acceptance or rejection of a host plant based on the presence or absence of stimulant (e.g., sugars, amino acids, vitamins) or deterrent chemicals, and moisture (Dethier, 1980; Schoonhoven, 1981; Städler, 1984; Otter, 1992; Krokos et al., 2002). Free amino acids, for example, elicit electrophysiological responses from the sensillae of lepidopteran larvae (Städler, 1984; Blaney & Simmonds, 1988). Many free essential amino acids (essential for insect growth and development) accumulate in plant tissues during water deficit stress in crop plants that range from cotton to sugarcane, Saccharum species, to pine trees, Pinus species (Mattson & Haack, 1987; Showler, 2012 ). Amino acids were even found to be more important determinants of corn susceptibility to neonate fall armyworms than toxins or other biochemical factors (Hedin et al., 1990). Resistance against the sugarcane aphid, Melanaphis sacchari (Zehnter), and the yellow sugarcane aphid, Sipha flava (Forbes), involved absence of some free essential amino acids in resistant sugarcane varieties (Akbar et al., 2010). Free amino acids are more available for use by herbivorous arthropods because insects absorb nitrogen through the gut mostly as free amino acids or small peptides (Brodbeck & Strong, 1987). Hence, enhanced foliar nutritional value as a result of water deficit is known to be an important determinant of neonate lepidopteran performance (Mattson, 1980; English-Loeb et al., 1997; Showler, 2001, 2012 ; Showler & Moran, 2003; Moran & Showler, 2005; Chen et al., 2008). In terms of water deficit stress, the mealybug Phenacoccus herreni Cox & Williams develops and reproduces better on drought stressed than on well watered cassava, Manihot esculenta Crantz, in response to greater concentrations and more nutritious combinations of free amino acids (Calatayud et al., 2002). The eldana borer, Eldana saccharina Walker, a stalkborer of sugarcane in Africa, prefers water deficit stressed host plants (Moyal, 1995), and the European corn borer, Ostrinia nubilalis (Hübner), inflicts up to twice the injury to water deficit stressed corn than to corn under conventional irrigation (Godfrey et al., 1991). Correlations were reported between elevated free amino acid concentrations in phloem sap of water deficit stressed wheat, Triticum aestivum L., and barley, Hordeum vulgare L., and population increases by the bird oat-cherry aphid (Weibull, 1987) and the cabbage aphid on Brassica spp. (Cole, 1997). Similarly, bark beetle outbreaks during times of drought are associated with greater concentrations of amino acids (and soluble sugars) in host plant phloem that likely contribute toward improved scolytid performance (Mattson & Haack, 1987).

In addition to elevated levels of free essential amino acids, free proline, a nonessential amino acid that accumulates in most water deficit-afflicted plants, is a feeding stimulant for many phytophagous arthropods (Mattson & Haack, 1987; Städler, 1984). Dadd (1985) reported that a number of amino acids, particularly glycine, alanine, serine, methionine, histidine, proline, and γ-aminobutyric acid, were phagostimulants to a number of insect species. Amino acids that elicited the greatest response as feeding stimulants to southwestern corn borer larvae were determined to be arginine, histidine, lysine, methionine, phenylanaline, valine (essentials), alanine, glycine, and serine (nonessentials) (Hedin et al., 1990), but not proline.

Water deficit stress has also been associated with increased concentrations of carbohydrates (which have important roles in osmotic adjustment) in many plants (Schubert et al., 1995; Kameli & Lösel, 1996; Massacci et al., 1996; Mohammadkhani & Heidari, 2008). Corn plants with elevated soluble carbohydrate concentrations were preferred by the European corn borer for oviposition (Derridj & Fiala, 1983; Derridj et al., 1986), and styloconic sensilla of larvae and adults of three noctuid species were highly responsive to sugars, especially sucrose and fructose (Blaney & Simmonds, 1988). These two sugars are known to be important feeding stimulants for both life stages (Frings & Frings, 1956; Blom, 1978), and fructose, glucose, maltose, and sucrose have been identified as phagostimulants for other insects (Bernays, 1985). Electrophysiological recordings revealed that the maxillary sensilla styloconica of fifth instar African armyworm, Spodoptera exempta (Walker), and the lepidopteran stalkborers E. saccharina, Maruca testulalis (Geyer), and Chilo partellus (Swinhoe), were stimulated by 13 different carbohydrates (Otter, 1992). In an experiment involving fall armyworm larval feeding, sucrose elicited ≥5-fold more feeding response than fructose or glucose (Hedin et al., 1990). Carbohydrates are well known as sources of energy for arthropods, and they are therefore highly important as nutrients (Nation, 2002). Studies on larval rice stem borers, for instance, showed that fructose, glucose, and sucrose are highly nutritious as compared with other carbohydrates based on their growth and development (Ishii et al., 1959; Ishii, 1971). Also, eastern spruce budworm, Choristoneura fumiferana Clemens, outbreaks often follow droughts (Mattson & Haack, 1987) because water deficit stressed trees accumulate sugar and sugar alcohols (Price, 2002).

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3. Water is a nutrient, too

Water deficit affects both the availability of water, which is a nutrient itself, to herbivores as well as the nutritional quality of dietary biochemical components that accumulate as osmoprotectants or for other purposes. When herbivorous arthropods are unable to have access to sufficient amounts of wager, their populations can decline. For example, aphid populations are reduced under conditions of continued and severe host plant water deficit ( Showler, 2012 ). Black bean aphid, Aphis fabae Scopali, survivorship was diminished on continuously drought stressed sugar beet, Beta vulgaris L., leaves (Kennedy & Booth, 1959), and reproduction and survival were negatively affected for the mustard aphid, Lipaphis erysimi (Kalt.) on radish, Raphanus sativus L. (Sidhu & Kaur, 1976); the spotted alfalfa aphid, Therioaphis maculata (Buckton), on alfalfa, Medicago sativa L. (McMurtry, 1962); the greenbug on sorghum, Sorghum bicolor (L.) Moench (Michels & Undersander, 1986); the potato aphid, Macrosiphum euphorbiae (Thomas), on potato, Solanum tuberosum L. (Nguyen et al., 2007); the bird oat-cherry aphid, Rhopalosiphum padi (L.), on tall fescue (Bultman & Bell, 2003); and the eastern spruce gall adelgid, Adelges abietis (L.), on Norway spruce, Picea abies (L.) Karst. (Bjőrkman, 2000). The most likely cause of the host plants’ unsuitability for aphids under such conditions is low turgor which reduces the ability of aphids to feed (Levitt, 1951; Wearing & Van Emden, 1967). Turgor facilitates aphid ingestion by forcing fluids out of the plant and through the aphids’ stylet lumens (Kennedy & Mittler, 1953; Maltais, 1962; Auclair, 1963: Magyarosy & Mittler, 1987; Douglas & Van Emden, 2007); turgor loss reduces or curtails feeding by aphids despite their cybarial pump. This has been reported to occur for the black bean aphid on different plant hosts (Kennedy et al., 1958); the cotton aphid, Aphis gossypii Glover on cotton, Gossypium hirsutum L. (Komazaki, 1982); the greenbug on wheat (Sumner et al., 1983); and the pea aphid, Acyrthosiphon pisum Harris, on alfalfa (Girousse & Bournoville, 1994). Also, greater concentrations of host plant osmolytes and other biochemicals associated with drought stress increase sap viscosity which resists flow through the stylets (Douglas & Van Emden, 2007), impeding ingestion despite the enriched nutritional quality of the sap (Kennedy et al., 1958).

The greater nutritional quality of water deficit stressed plants can be offset by the condition that causes it: insufficient water. When provided with dried, ground material from water-deficit stressed tomato plants, Lycopersicon esculentum Mill., incorporated into a nonnutritive diet, beet armyworm, Spodoptera exigua (Hübner), larval growth decreased (English-Loeb et al., 1997). Cecropia moth, Hyalophora cecropia L., larvae reared on water deficit stressed wild cherry, Prunus serotina Ehrh., leaves grew more slowly than those fed on well-watered plants, but they, and beet armyworm larvae on water deficit stressed cotton leaves, consumed greater quantities of leaf tissue in order to gain access to more water, and possibly in order to supplement body water with water derived from respiration (Scriber, 1977; Showler & Moran, 2003 ). Under field conditions, fall armyworm; soybean looper, Pseudoplusia includens (Walker); and beet armyworm larval survivorships increased and development was hastened in soybeans that were irrigated compared with dryland-grown soybeans (Huffman & Mueller, 1983). These observations suggest that soft-bodied lepidopteran larvae that live on plant surfaces exposed to the desiccating effects of direct sunlight and ambient air (unlike lepidopteran stalkboring larvae that live in moist plant interiors) are especially vulnerable to the desiccating effects of insufficient water supply.

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4. Some non-nutrient-related associations of water deficit with phytophagous arthropods

Host plant selection among insects also involves visual and physical factors such as leaf shape, color, and size (Ramaswamy, 1988; Renwick & Radke, 1988; Renwick & Chew, 1994; Showler & Castro, 2010b), and both constitutive and inducible plant chemical defenses can vary in response to water deficit stress (Lombardero et al., 2000), but visual and physical cues, and defensive compounds are not considered as being nutritional for the purposes of this chapter (although defensive compounds might loosely be considered as being types of nutrients, they mostly repel, interfere with feeding, or act as toxins). Concentrations of several classes of defensive secondary compounds tend to increase in plant tissues in response to moderate drought, including terpenoids (some of which are attractants (Mattson & Haack, 1987) and alkaloids (Gershenson, 1984; Hoffmann et al., 1984; Sharpe et al., 1985; Lorio, 1986; Mattson & Haack, 1987; Showler, 2012 ), but intensified drought stress can lead to reductions of these compounds (Mattson and Haack, 1987). Drought can also influence predator and parasitoid guilds that affect phytophagous arthropod populations ( Showler, 2012 ), but plant stress is not directly involved. Other mechanisms that might also contribute toward plant vulnerability to herbivorous arthropods under conditions of water deficit stress have been suggested (Mattson & Haack, 1987), including acoustical cues, detoxification of foods by drought stressed insects, and drought-induced genetic changes in arthropods, but they have not been well substantiated.

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5. Multiple effects of water deficit: case study on sugarcane and the Mexican rice borer

The Mexican rice borer, Eoreuma loftini (Dyar), and its association with sugarcane is arguably one of the most illustrative examples of how an economically important phytophagous arthropod is affected by limited availability of water. The crambid moth is indigenous to western Mexico (Morrill, 1925; Van Zwaluwenberg, 1926) where it is a major pest of sugarcane, but it had spread by the mid 1970s to Veracruz, San Luis Potosi, and Tamaulipas in eastern Mexico (Johnson, 1984). First detected in the United States in the Lower Rio Grande Valley of Texas in 1980 ( Johnson, 1981 , 1984; Johnson & Van Leerdam, 1981 ), the pest dispersed into rice producing areas of east Texas (Browning et al., 1989; Reay-Jones et al., 2008), and in 2008 it moved into Louisiana (Hummel et al., 2008, 2010). Because the Mexican rice borer was recently determined to prefer corn over other crop plants (Showler et al., 2011), its assumed range might be considerably underestimated ( Showler & Reagan, 2012 ).

Eggs are mostly deposited in clusters within folds of dry sugarcane leaves, although eggs are also laid in folded green living tissue if available (Showler & Castro, 2010b). Van Leerdam et al. (1986) found 96% of the pest’s eggs on the basal 80 cm of sugarcane plants where most dry leaf tissue is located. The Mexican rice borer is not so much stress-oriented as it is nutritionally-oriented in that it prefers to lay eggs on dry foliage of plants stressed by limited water and of plants growing in enriched soil (Showler & Castro, 2010a; Showler & Reagan, 2012). Water deficit stress in sugarcane plants, however, unlike over-fertilized plants, offers increased quantities of dry, folded leaf tissue per plant, contributing to the crop’s vulnerability (Reay-Jones et al., 2005; Showler & Castro, 2010b). In a greenhouse no-choice cage experiment using sugarcane plants from which all dry leaf tissue was excised and removed from the cages, or placed at the bottom of the cages like a mulch, and intact (dry leaf tissue remained on the plants) sugarcane plants (controls), numbers of eggs and the degree of larval infestation was distinctly greater on the controls (Figs. 1 & 2; Showler & Castro, 2010b).

Figure 1.

Mean (± SE) numbers of Mexican rice borer eggs on green and dry leaf tissue per sugarcane plant; ANOVA, Tukeys HSD (P < 0.05), n = 7 replicates per assay (Showler & Castro, 2010b).

Early instars feed on living leaf tissue, under fresh leaf sheaths, and some tunnel into the leaf midrib; later instars bore into the main stalk (Wilson, 2011). Injury from stalk tunneling results in deadheart, decreased sugar production, and stunting or lodging of stalks sometimes so severe that harvest becomes unfeasible (Johnson, 1985; Legaspi et al., 1997; Hummel et al., 2008). Tunnels within host plant stalks are packed with frass, blocking entry of predators and parasitoids (Hummel et al., 2008). Pupation occurs within the stalk after mature larvae make emergence holes protected with a thin window of outer plant tissue (Hummel et al., 2008). In the Lower Rio Grande Valley, a life cycle takes 30–45 days, and there are 4–6 overlapping generations per year (Johnson, 1985; Legaspi et al., 1997). Tunneling damage and the insect’s prevalence has made it the key sugarcane pest of south Texas, displacing the sugarcane borer, Diatraea saccharalis (F.) (Van Leerdam et al., 1984; Legaspi et al., 1997).

Figure 2.

Mean (± SE) numbers of Mexican rice borer larval entry holes per sugarcane stalk; ANOVA, Tukeys HSD (P < 0.05), n = 7 replicates per assay (Showler & Castro, 2010b).

Approximately 20% of sugarcane internodes are injured by Mexican rice borers in south Texas, and larval entry holes also provide portals for red rot, resulting in additional loss of sugar (Van Zwaluwenberg, 1926; Osborn & Phillips, 1946; Johnson, 1985). On some varieties of sugarcane, up to 50% bored internodes have been reported ( Johnson, 1981 ); Mexican rice borer injury results in losses of US$575 per hectare of sugarcane (Meagher et al., 1994) and US$10–20 million annually (Legaspi et al., 1997, 1999). Projected economic consequences of Mexican rice borer infestation of Louisiana includes US$220 million in sugarcane and US$45 million in rice (Reay-Jones et al., 2008). In corn, stalk boring and secondary infection by stalk rot pathogens can cause shattering, lodging, and complete collapse of stalks (Showler et al., 2011) such that by season’s end >50% of stalks of susceptible varieties are destroyed (Showler, unpublished data).

A connection between irrigation practices and severity of Mexican rice borer infestation was first suggested by Meagher et al. (1993), and later studies indicated that drought stressed sugarcane is preferred for oviposition because there is more dry leaf tissue and the nutritional value, at least in terms of a number of important free amino acids, is enhanced (Tables 1 & 2) (Muquing & Ru-Kai, 1998; Reay-Jones et al., 2005, 2007; Showler & Castro, 2010a). Although severe water deficit stress of sugarcane reduces sugar production, some cultivars under moderate stress accumulate sugars (Hemaprabha et al., 2004), and Mexican rice borer preference among species of host plants (Showler et al., 2011) has been associated with concentrations of fructose (Showler, unpublished data). Differences in oviposition preference were not observed on excised dry leaf tissue regardless of whether the sugarcane plant from which it originated was water deficit stressed or well watered; hence, the expression of sugarcane vulnerability or resistance appears to require the pest’s ability to detect nutrients in living leaf tissue (Showler & Castro, 2010b). Although a sugarcane cultivar with some degree of resistance to the Mexican rice borer was still better protected than a susceptible variety under drought conditions, water deficit increased injury to the crop by ≈2.5-fold in each (Reay-Jones et al., 2005). Reay-Jones et al. (2003) also reported that high soil salinity, a stress factor that also heightens free amino acid accumulations in plants (Labanauskas et al., 1981; Cusido et al., 1987), increases Mexican rice borer infestations in sugarcane. Further, relatively high concentrations of organic matter incorporated into soil of the Lower Rio Grande Valley (and conventionally fertilized with nitrogen) resulted in 18% more stalk production per sugarcane stool but this effect was offset by substantial increases in Mexican rice borer infestation, causing stalk weight, length, and percentage brix reductions relative to sugarcane fertilized with conventional nitrogen fertilizer or chicken litter (Showler, unpublished data). The composted soil was associated with greater accumulations of free amino acids and fructose (Showler, unpublished data). These associations reveal that the pest is not responding simply to water deficit, but instead to nutritional enhancement of the plant whether moderated by stress or by other factors.

Table 1.

Mean (± SE) water potential (bar), and numbers of dry leaves, Mexican rice borer egg clusters, total eggs, entry holes, and exit holes per stalk of two sugarcane varieties maintained under well watered or drought stressed greenhouse conditions (Showler & Castro, 2010a)

a One-way ANOVA, randomized complete block design, df = 3,15.


b W, well watered; D, drought stressed.


Table 2.

Mean (± SE) picomoles of free amino acid per μl of sugarcane leaf juice in two varieties, L97-128 and CP70-321, that were well watered or drought stressed (Showler & Castro, 2010a)

Means within each row followed by different letters are significantly different (P < 0.05).


a Cystine was detectable but not found in the samples.


b One-way ANOVA, randomized complete block design, df = 3, 12.


c W, well watered; D, drought stressed


In addition to water deficit stress associations with Mexican rice borer preferences for physical (i.e., dry, curled leaf tissue) and nutritional factors (i.e., amino acids and possibly sugar accumulations), water availability has a strong influence on abundances of a voracious predator, the red imported fire ant, Solenopsis invicta Buren, which has already been shown to be an efficient predator of the stalk boring moth, D. saccharalis, in Louisiana ( Showler, 2012 ; Showler & Reagan, 2012 ). Originally from wet habitats of South America, the red imported fire ant entered the United States in 1929 and it spread throughout much of the wet southern states (Lofgren, 1986). To provide another example of the predator’s effectiveness against insect pests, red imported fire ant foraging activity accounts for 58% of boll weevil mortality along the relatively wet coastal cotton-growing region of Texas (Sturm & Sterling, 1990), and red imported fire ant predation on immature boll weevils averaged 84% compared with 0.14% and 6.9% mortality caused by parasitism and desiccation, respectively (Fillman & Sterling, 1983). In the drier subtropics of south Texas, however, even in cotton with rank weed growth commonly associated with thriving red imported fire ant populations in wetter regions (Showler et al., 1989; Showler & Reagan, 1991), few or no red imported fire ants were found and boll weevil infestations were not affected by predation ( Showler & Greenberg, 2003 ). While sugarcane in relatively dry regions, such as south Texas, is not protected by red imported fire ants, it is possible that the predator’s greater abundance in the more moist sugarcane growing conditions of Louisiana will suppress Mexican rice borer populations ( Showler & Reagan, 2012 ) despite its cryptic larval behavior.

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6. Conclusion

Water deficit might initially appear to affect herbivorous arthropod populations because of a single factor, but the associations of the Mexican rice borer with water indicate a more complex relationship that can involve physical, biochemical, and ecological factors. Levels of Mexican rice borer infestation are likely influenced by low water availability in at least three ways, only one of which is directly related to the nutritional status of the crop. Drought changes many environmental conditions relative to arthropods, such as soil condition, leaf size and color, lignification of plant cell walls, secondary protective compounds, and natural enemy activity, but accumulations of nutrients, particularly free amino acids and carbohydrates, unlike the other drought-related conditions, directly result from water deficit stress to the plant. This plant stress response to water deficit influences levels of pest infestations by causing the plant emit volatile semiochemicals and by enhancing the nutritional quality of the plant. Water deficit can also make it difficult for some plant sucking insects (e.g., aphids) to attain water and nutrients, and soft-bodied lepidopteran larvae living on surfaces of water deficit stressed plants ingest insufficient amounts of water to sustain themselves against desiccation despite compensating by consuming greater quantities of plant tissue. While non-nutritional factors are often important under conditions of water deficit, the nutritional status of the plant to herbivorous arthropods is directly modulated by water deficit stress, and host plant nutritional quality is arguably the most fundamental component of plant-herbivore interactions.

References

  1. 1. Akbar W. Showler A. T. White W. H. Reagan T. E. 2010Categorizing sugarcane cultivars for resistance to the sugarcane aphid and yellow sugarcane aphid (Hemiptera: Aphididae). Journal of Economic Entomology 103 1431 1437 0022-0493
  2. 2. Apelbaum A. Tang S. F. 1981 Biosynthesis of stress ethylene induced by water deficit Plant Physiology 68 594 596 0981-9428
  3. 3. Archer T. L. Bynum E. D. Jr Onken A. B. Wendt C. W. 1995Influence of water and nitrogen fertilizer on biology of the Russian wheat aphid (Homoptra: Aphdidae) on wheat. Crop Protection 14 165 169 0261-2194
  4. 4. Assefa Y. Conlong D. E. Van den Berg. J. Le Rü B. P. 2008The wider distribution of Eldana saccharina (Lepidoptera: Pyralidae) in South Africa and its potential risk to maize production. Proceedings of the South African Sugarcane Technologists Association 81 290 297 0370-1816
  5. 5. Auclair J. L. 1963 Aphid feeding and nutrition. Annual Review of Entomology 8 439 491 0066-4170
  6. 6. Bailey W. 2000Notes from Missouri’s fields. Integrated Pest and Crop Management Newsletter 10 1 2 0838-0937
  7. 7. Becana M. Moran J. F. Iturbe-Ormaete I. 1998 Iron-dependent oxygen free radical generation in plants subjected to environmental stress: toxicity and antioxidant protection Plant and Soil 201 137 147 0003-2079X.
  8. 8. Beck E. H. Fettig S. Knake C. Hartig K. Bhattarai T. 2007 Specific and unspecific responses of plants to cold and drought stress Journal of Bioscience 32 501 510 0973-7138
  9. 9. Bernays E. A. 1985Regulation of feeding behavior. In Insect Physiology, Biochemistry, and Pharmacology: Regulation, Digestion, Nutrition, Excretion, G.A. Kerbut, G.A. & Gilbert, L.I. (eds.), 1 32Pergamon Press, 100080308058New York, USA.
  10. 10. Bernays E. A. Chapman R. F. 1994 Host-Plant Selection by Phytophagous Insects Chapman and Hall, 100412031310York, New York, USA.
  11. 11. Bjőrkman C. 2000Interactive effects of host resistance and drought stress on the performance of a gall-making aphid living on Norway spruce. Oecologia 123 223 231 0029-8549
  12. 12. Blaney W. M. Simmonds M. S. J. 1988Food selection in adult and larvae of three species of Lepidoptera: a behavioral and electrophysiological study. Entomologia Experimentalis et Applicata 49 111 121 0013-8703
  13. 13. Blom F. 1978 Sensory activity and food intake: a study of input-output relationships in two phytophagous insects.Nederlands Journal of Zoology 28 277 340 0028-2960
  14. 14. Boyer J. S. 1982 Plant productivity and environment. Science 218 443 448 0036-8075
  15. 15. Branco M. Pereira J. S. Mateus E. Tavares C. Paiva M. R. 2010 Water stress affects Tomicus destruens host pine preference and performance during the shoot feeding phase Annals of Forest Science doi:forest/201021, 1286-4560
  16. 16. Brodbeck B. Strong D. 1987Amino acid nutrition of herbivorous insects and stress to host plants. In Insect Outbreaks; Barbosa, P. & Schultz, J. C. (eds.), 346 364Academic Press, 0-12078-148-4England.
  17. 17. Browning H. W. Way M. O. Drees B. M. 1989 Managing the Mexican rice borer in TexasTexas Agricultural Extension Service Bulletin B-1620.
  18. 18. Bultman T. L. Bell G. D. 2003Interaction between fungal endophytes and environmental stress influences plant resistance to insects. Oikos 103 182 190 0030-1299
  19. 19. Burgess A. J. Warrington S. Allen-Williams L. 1994Cabbage aphid (Brevicoryne brassicae L.) ‘performance’ on oilseed rape (Brassica napus L.) experiencing water deficiency: roles of temperature and food quality. Acta Horticulturae 407: ISHS Brassica Symposium- IX Crucifer Genetics Workshop, 16 19 0567-7572
  20. 20. Burton R. L. Schuster D. J. 1981Oviposition stimulant for tomato pinworm from surfaces of tomato plants. Annals of the Entomological Society of America 74 512 515 0013-8746
  21. 21. Bussis D. Heineke D. 1998 Acclimation of potato plants to polyethylene glycol-induced water deficit. II. Contents and subcellular distribution of organic solutes. Journal of Experimental Botany 49 1361 1370 1460-2431
  22. 22. Calatayud P. A. Polania M. A. Seligmann C. D. Bellotti A. C. 2002 Influence of water-stressed cassava on Phenacoccus herreni and three associated parasitoids Entomologia Experimentalis et Applicata 102 163 175 0013-8703
  23. 23. Cates R. G. Redack R. A. 1988Variation in the terpene chemistry of Douglas fir and its relationship to western spruce budworm success. In Chemical Mediation of Coevolution; Spencer, K.C. (ed.), 317 344Academic Press, 978-0-12656-856-1San Diego, California, USA.
  24. 24. Chen Y. Ruberson J. R. Olson D. M. 2008 Nitrogen fertilization rate affects feeding, larval performance, and oviposition preference of the beet armyworm, Spodopteraexigua, on cotton. Entomologia Experimentalis et Applicata 126 244 255 0013-8703
  25. 25. Chrominsky A. Visscher-Neumann S. Jurenka R. 1982Exposure to ethylene changes nymphal growth rate and female longevity in the grasshopper Melanaphis sanguinipes. Naturwissenschaften 69 45 66 0028-1042
  26. 26. Cole R. A. 1997 The relative importance of glucosinolates and amino acids to the development of two aphid pests Brevicoryne brassicae and Myzus persicae on wild and cultivated brassica species Entomologia Experimentalis et Applicata 85 121 133 0013-8703
  27. 27. Cusido R. M. Palazon J. Altabella T. Morales C. 1987 Effect of salinity on soluble protein, free amino acids and nicotine contents in Nicotiana rustica L Plant and Soil 102 55 60 0003-2079X.
  28. 28. Dadd R. H. 1985Nutrition: organisms. In Comprehensive Insect Physiology, Biochemistry and Pharmacology: Regulation, Digestion, Nutrition, Excretion, Kerkut, G.A. & Gilbert, L.I. (eds.), 315 319Pergammon Press, 100080308058New York, USA.
  29. 29. Derridj S. Fiala V. 1983Sucres solubles des feuilles de mais (Zea mays L.) et oviposition de la pyrale (Ostrinia nubilalis Hbn.). Comptes Rendues Academie Agriculture Francais 69 465 472 0001-3986
  30. 30. Derridj S. Fiala V. Jolivet E. 1986 Increase of European corn borer (Ostrinia nubilalis) oviposition induced by a treatment of maize plants with maleic hydrazide: role of leaf carbohydrate. Entomologia Experimentalis et Applicata 41 305 310 0013-8703
  31. 31. Dethier V. G. 1980 Evolution of receptor sensitivity to secondary plant substances with special reference to deterrents American Naturalist 115 45 66 0003-0147
  32. 32. Dix M. E. Cunningham A. A. King R. M. 1996 Evaluating spring cankerworm (Lepidoptera: Geometridae) preference for Siberian elm clones Environmental Entomology 25 58 62 0004-6225X.
  33. 33. Dorschner K. W. Johnson R. C. Eikenbary R. D. Ryan J. D. 1986Insect-plant interactions: greenbug (Homoptera: Aphididae) disrupt acclimation of winter wheat to drought stress. Environmental Entomology 15 118 121 0004-6225X.
  34. 34. Douglas A. E. Van Emden H. F. 2007Nutrition and symbiotes. In Aphids as Crop Pests Van Emden, H.F. & Harrington, R., (eds.), 112 134CAB International, 1-96399-939-6Park, England.
  35. 35. Dubey R. S. 1999Protein synthesis by plants under stressful conditions. In Handbook of Plant and Crop Stress, Pessarakli, M. (ed.), 365 386Marcel Dekker, 978-1-43981-396-6New York, New York, USA.
  36. 36. Dunn J. P. Kimmerer T. W. Nordin G. L. 1986 Attraction of the twolined chestnut borer, Agrilus bilineatus (Weber) (Coleoptera: Buprestidae), and associated borers to volatiles ofstressed white oak. Canadian Entomologist 118 503 509
  37. 37. Ebel R. C. Mattheis J. P. Buchanan D. A. 1995 Drought stress of apple trees alters leaf emissions of volatile compounds Physiologia Plantarum 92 709 712 0031-9317
  38. 38. English-Loeb G. Stout M. J. Duffey S. S. 1997Drought stress in tomatoes: changes in plant chemistry and potential nonlinear consequences for insect herbivores. Oikos 79 456 468 0030-1299
  39. 39. Fan L. Linker R. Gepstein S. Tanimoto E. Yamamoto R. Neumann P. M. 2006Progressive inhibition by water deficit of cell wall extensibility and growth along the elongation zone of maize roots is related to increased lignin metabolism and progressive stellar accumulation of wall phenolics. Plant Physiology 140 603 612 0981-9428
  40. 40. Fenemore P. G. 1980Oviposition of potato tuber moth, Phthorimaea operculella Zell. (Lepidoptera: Gelechiidae): identification of host-plant factors influencing oviposition response. New Zealand Journal of Zoology 7 435 439 0301-4223
  41. 41. Fillman D. A. Sterling W. L. 1983Killing power of the red imported fire ant (Hym.: Formicidae): a key predator of the boll weevil (Coleoptera: Curculionidae). Entomophaga 28 339 344 0013-8959
  42. 42. Frings H. Frings M. 1956The loci of contact chemoreceptors involving in feeding reactions in certain Lepidoptera. Biology Bulletin 110 291 199 0006-3185
  43. 43. Garg B. K. Kathju S. Burman U. 2001Influence of water stress on water relations, photosynthetic parameters and nitrogen metabolism of moth bean genotypes. Biologia Plant 44 289 292 1573-8264
  44. 44. Gershenson J. 1984Changes in the levels of plant secondary metabolites under water and nutrient stress. Recent Advances in Phytochemistry 18 273 320 0079-9920
  45. 45. Girousse C. Bournoville R. 1994Role of phloem sap quality characteristics on performance of pea aphid grown on lucerne genotypes. Entomologia Experimentalis et Applicata 70 227 235 0013-8703
  46. 46. Godfrey L. D. Holtzer T. O. Spomer S. M. Norman J. M. 1991European corn borer (Lepidoptera:: Pyralidae) tunneling and drought stress: effects on corn yield. Journal of Economic Entomology 54 1850 1860 0022-0493
  47. 47. Golan-Goldhirsch A. N. Samish S. Agami M. Lips H. 1989The relationship between some perennial desert plants originated in different phytogeographical regions and proline concentration. Journal of Arid Environments 17 327 333 0140-1963
  48. 48. Haack R. A. Slansky F. Jr 1987Nutritional ecology of wood-feeding Coleoptera, Lepidoptera, and Hymenoptera. In The Nutritional Ecology of Insects, Mites, and Spiders; Slansky, F., Jr. & Rodriguez, J.G. (eds.), 449 486John Wiley & Sons, 10New York, New York, USA.
  49. 49. Hansen A. D. Hitz W. D. 1982Metabolic responses of mesophytes to plant water stress. Annual Review of Plant Physiology 33 163 203 0079-9920
  50. 50. Harfouche A. L. Shivaji R. Stocker R. Williams P. W. Luthe D. S. 2006Ethylene signaling mediates a maize defense response to insect herbivory. Molecular Plant-Microbe Interactions 19 189 199 0094-0282
  51. 51. Hedin P. A. Thompson A. C. Gueldner R. C. 1976Cotton plant and insect constituents that control boll weevil behavior and development. In Biochemical Interactions Between Plants and Insects, Wallace, J.W. & Mansek, R.K. (eds.), 271 350Plenum, 0-30634-710-5York, New York, USA.
  52. 52. Hedin P. A. Davis F. M. Williams W. P. Salin M. L. 1984Possible factors of leaf-feeding resistance in corn to the southwestern corn borer. Journal of Agricultural and Food Chemistry 32 262 267 0021-8561
  53. 53. Hedin P. A. Williams W. P. Davis F. M. Buckley P. M. 1990Roles of amino acids, protein, and fiber in leaf-feeding resistance of corn to the fall armyworm. Journal of Chemical Ecology 16 1977 1995 0098-0331
  54. 54. Hemaprabha G. Nagarajan R. Alarmelu S. 2004Responses of sugarcane genotypes to water deficit stress. Sugar Tech 6 165 168 0972-1525
  55. 55. Hodges J. D. Lorio P. L. Jr 1975Moisture stress and composition of xylem oleoresin in loblolly pine. Forest Science 21 283 290 0001-5749X.
  56. 56. Hoffman J. J. Kingsolver B. E. Mc Laughlin S. P. Timmermann B. N. 1984Production of resins by arid-adapted Asteraea. Recent Advances in Phytochemistry 18 251 271 0079-9920
  57. 57. Hsiao T. C. 1973Plant responses to water stress. Annual Review of Plant Physiology 24 519 570 0079-9920
  58. 58. Huffman F. R. Mueller A. J. 1983Effects of beet armyworm (Lepidoptera: Noctuidae) infestation levels on soybean. Journal of Economic Entomology 76 744 747 0022-0493
  59. 59. Hummel N. Reagan T. E. Pollet D. Akbar W. Beuzelin J. M. Carlton C. Saichuk J. Hardy T. Way M. O. 2008Mexican rice borer, Eoreuma loftini (Dyar). Louisiana State University AgCenter Pub. 3098, Baton Rouge, LA.
  60. 60. Hummel N. A. Hardy T. Reagan T. E. Pollet D. Carlton C. Stout M. J. Beuzelin J. M. Akbar W. White W. H. 2010Monitoring and first discovery of the Mexican rice borer, Eoreuma loftini (Lepidoptera: Crambidae) in Louisiana. Florida Entomologist 93 123 124 0015-4040
  61. 61. Ingram J. Bartels D. 1996The molecular basis of dehydration tolerance in plants. Annual Review of Plant Physiology and Plant Molecular Biology 47 377 403 1040-2519
  62. 62. Ishii S. Azim A. Hirano C. 1959A further experiment on the effect of dietary levels of protein and carbohydrate on the growth of the rice stem borer, Chilo suppressalis larvae. Japanese Journal of Applied Entomology and Zoology 3 143 145 0021-4914
  63. 63. Ishii S. 1971Nutritional studies of the rice stem borer, Chilo suppressalis Walker, and its mass rearing. Entomophaga 16 165 173 0013-8959
  64. 64. Janagouar B. S. Venkatasubbaiah D. Janardhan K. V. Panchal Y. C. 1983Effect of short term stress on free proline accumulation, relative water content and potassium content in different plant parts of three cotton genotypes. Indian Journal of Plant Physiology 26 82 87 0019-5502
  65. 65. Johnson K. J. R. 1981 Acigona loftini (Lepidoptera: Pyralidae) in the Lower Rio Grande Valley of Texas, 1980-1981. In Proceedings of the 2 nd Inter-American Sugar Cane Seminar (Insect and Rodent Pests), Miami, Florida, USA, 166 171
  66. 66. Johnson K. J. R. 1984Identification of Eoreuma loftini (Dyar) (Lepidoptera: Pyralidae) in Texas, 1980: forerunner for other sugarcane boring pest immigrants from Mexico? Bulletin of the Entomological Society of America 30 47 52 1046-2821
  67. 67. Johnson K. J. R. 1985Seasonal occurrence and insecticidal suppression of Eoreuma loftini (Lepidoptera: Pyralidae) in sugarcane. Journal of Economic Entomology 78 960 966 0022-0493
  68. 68. Johnson K. J. R. Van Leerdam M. B. 1981Range extension of Acigona loftini into the Lower Rio Grande Valley of Texas. Sugar y Azucar 76: 119, 0039-4742
  69. 69. Jones C. G. 1991Plant stress and insect herbivory: toward an integrated perspective. In Responses of Plants to Multiple Stresses, Mooney, H.A.: Winner, W.E. & Pell, E.J. (eds.), 249 280Academic Press, 978-0-12505-355-6New York, New York, USA.
  70. 70. Jordan W. R. Ritchie J. T. 1971Influence of soil water stress on evaporation, root absorption, and internal water status of cotton. Plant Physiology 48 783 788 0981-9428
  71. 71. Kameli A. Lösel D. M. 1996Growth and sugar accumulation in durum wheat plants under water stress. New Phytology 132 57 62 1469-8137
  72. 72. Kennedy J. S. Mittler T. E. 1953A method for obtaining phloem sap via the mouth-parts of aphids. Nature 171: 528, 0028-0836
  73. 73. Kennedy J. S. Booth C. O. 1959Responses of Aphis fabae Scop. To water shortage in host plants in the field. Entomologia Experimentalis et Applicata 2 1 11 0013-8703
  74. 74. Kennedy J. S. Lamb K. P. Booth C. O. 1958Responses of Aphis fabae Scop. To water shortage in host plants in pots. Entomologia Experimentalis et Applicata 1 274 290 0013-8703
  75. 75. Kimmerer T. W. Kozlowski T. T. 1982Ethylene, ethane, acetaldehyde, and ethanol production by plants under stress. Plant Physiology 69 840 847 0981-9428
  76. 76. Klubertanz T. H. Pedigo L. P. Carlson R. E. 1990Effects of plant moisture stress and rainfall on population dynamics of the twospotted spider mite (Acari: Tetranychidae). Environmental Entomology 19 1773 1779 0004-6225X.
  77. 77. Knight H. Knight M. R. 2001Abiotic stress signaling pathways: specificity and cross-talk. Trends in Plant Science 6 262 267 1360-1385
  78. 78. Komazaki S. 1982Effects of constant temperature on population growth of three aphid species, Toxoptera citricidies (Kirkaldy), Aphis citricola van der Goot, and Aphis gossypii Glover (Homoptera: Aphididae) on citrus. Applied Entomology and Zoology 17 75 81 0134-7605X.
  79. 79. Krokos F. D. Konstantopoulou M. A. Mazomenos B. E. 2002Chemical characterization of corn plant compounds by different extraction techniques and the role of potent chemicals in the reproductive behavior of the corn stalk borer, Sesamia nonagrioides. International Organization for Biological Control West Palearctic Region Section Bulletin 25 1 9 1049-9694
  80. 80. Labanauskas C. K. Stolzy L. H. Handy M. F. 1981Protein and free amino acids in field-grown cowpea seeds as affected by water stress at various growth stages. Plant and Soil 63 355 368 0003-2079X.
  81. 81. Legaspi J. C. Legaspi B. C. Jr Irvine J. E. Saldana R. R. 1997Mexican rice borer, Eoreuma loftini (Lepidoptera: Pyralidae) in the Lower Rio Grande Valley of Texas: its history and control. Subtropical Plant Science 49 53 64 1009-7791
  82. 82. Legaspi J. C. Legaspi B. C. Jr Irvine J. E. Meagher R. L. Jr Rozeff N. 1999Stalkborer damage on yield and quality of sugarcane in the Lower Rio Grande Valley of Texas. Journal of Economic Entomology 92 228 234 0022-0493
  83. 83. Levitt J. 1951Frost, drought and heat resistance. Annual Review of Plant Physiology 2 245 268 0079-9920
  84. 84. Lofgren C. S. 1986History of imported fire ants in the United States. In Fire Ants and Leaf-cutting Ants: Biology and Management, Lofgren, C.S. (ed.), 36 47Westview Press, 10Boulder, Colorado, USA.
  85. 85. Lombardero M. J. Ayres M. P. Lorio P. L. Jr Ruel J. J. 2000Environmental effects on constitutive and inducible resin defences of Pinus taeda. Ecology Letters 3 329 339 1461-0248
  86. 86. Lorio P. L. 1986Growth-differentiation balance: a basis for understanding southern pine beetle-tree interactions. Forest Ecology and Management 14 259 273
  87. 87. Lorio P. L. Jr Stephen F. M. Paine T. D. 1995Environment and ontogeny modify loblolly pine response to induced acute water deficits and bark beetle attack. Forest Ecology and Management 73 97 110
  88. 88. Magyarosy A. C. Mittler T. E. 1987Aphid feeding rates on healthy and beet curly top virus-infected plants. Phytoparasitica 15 335 338 0334-2123
  89. 89. Maltais J. B. 1962A simple apparatus for feeding aphids aseptically on chemically defined diets. Canadian Entomologist 84 291 294 0000-8347X.
  90. 90. Massacci A. Battistelli A. Loreto F. 1996Effect of drought stress on photosynthetic characteristics, growth and sugar accumulation of field-frown sweet sorghum. Australian Journal of Plant Physiology 23 331 340 0320-7844
  91. 91. Mattson W. J. J. 1980Herbivory in relation to plant nitrogen content. Annual Review of Ecology and Systematics 11 119 161 0066-4162
  92. 92. Mattson W. J. Haack R. A. 1987The role of drought in outbreaks of plant-eating insects. Bioscience 37 110 118 0006-3568
  93. 93. Mc Murtry J. A. 1962Resistance of alfalfa to spotted alfalfa aphid in relation to environmental factors. Hilgardia 32 501 539 0073-2230
  94. 94. Meagher R. L. Jr Pfannenstiel R. S. Saldana R. R. 1993Survey and estimated injury of the Mexican rice borer in Texas sugarcane. Journal of American Sugar Cane Technologists 13 22 26 0003-1216
  95. 95. Meagher R. L. Jr Smith J. W. Jr Johnson K. J. R. 1994Insecticidal management of Eoreuma loftini (Lepidoptera: Pyralidae) on Texas sugarcane: a critical review. Journal of Economic Entomology 87 1332 1344 0022-0493
  96. 96. Michels G. J. Jr Undersander D. J. 1986Temporal and spatial distribution of the greenbug (Homoptera: Aphididae) on sorghum in relation to water stress. Journal of Economic Entomology 79 1221 1225 0022-0493
  97. 97. Miller J. R. Strickler K. L. 1984Finding and accepting host plants. In Chemical Ecology of Insects, Bell, W. (ed.), 127 157Sinauer, 0-87893-070-1Massachusetts, USA.
  98. 98. Mohammadkhani N. Heidari R. 2008Drought-induced accumulation of soluble sugars and proline in two maize varieties. World Applied Sciences Journal 3 448 453 1818-4952
  99. 99. Moran P. J. Showler A. T. 2005Plant responses to water deficit and shade stresses in pigweed and their influence on feeding and oviposition by the beet armyworm (Lepidoptera: Noctuidae). Environmental Entomology 34 929 937 0004-6225X.
  100. 100. Morrill A. W. 1925Commercial entomology on the west coast of Mexico. Journal of Economic Entomology 18 707 716 0022-0493
  101. 101. Moyal P. 1995Borer infestation and damage in relation to the maize stand density and water stress in the Ivory Coast. International Journal of Pest Management 41 114 121 0967-0874
  102. 102. Muquing R. . Ku-Rai C. 1998Osmotic adjustment in leaves of sugarcane in response to water stress. Sugar Cane 5 3 7 0378-3774
  103. 103. Nation J. L. 2002Insect Physiology and Biochemistry. CRC Press, 101420061771Raton, Florida, USA.
  104. 104. Nguyen T. T. A. Michaud D. Cloutier C. 2007Proteomic profiling of aphid Macrosiphum euphorbiae responses to host-plant-mediated stress induced by defoliation and water deficit. Journal of Insect Physiology 53 601 611 0022-1910
  105. 105. Ormeño E. Mévy J. P. Vila B. Bousquet-Mélou A. Greff S. Bonin G. Fernandez C. 2007Water deficit stress induces different monoterpene and sesquiterpene emission changes in Mediterranean species. Relationship between terpene emissions and plant water potential. Chemosphere 67 276 284 0045-6535
  106. 106. Osborn H. T. Phillips G. R. 1946 Chilo loftini in California, Arizona, and Mexico. Journal of Economic Entomology 39 755 759 0022-0493
  107. 107. Otter C. J. den 1992Responses of the African armyworm and three species of borers to carbohydrates and phenolic substances: an electro- and behavioral physiological study. Entomologia Experimentalis et Applicata 63 27 37 0013-8703
  108. 108. Popov C. Trotus E. Vasilescu S. Barbulescu A. Rasnoveanu L. 2006Drought effect on pest attack in field crops. Romanian Agricultural Research 23 43 52 1222-4227
  109. 109. Price J. T. 2002Climate change, birds, and ecosystems- why should we care? In Managing for Healthy Ecosystems, Rapport, D.J.; Lasley, C.L.; Rolston, D.E.; Nielson, N.O.; Qualset, C.O. & Damania, A.B. (eds.), 60 61CRC Press, 978-1-42003-213-0Boca Raton, Florida, USA.
  110. 110. Ramanulu S. Kaiser W. Deitz K. J. 1999Salt and drought stress differentially affect the accumulation of extracellular proteins in barley. Journal of Bioscience 54 337 347
  111. 111. Ramaswamy S. B. 1988Host finding by moths: sensory modalities and behaviors. Journal of Insect Physiology 34 235 249 0022-1910
  112. 112. Ramos P. Rosales R. Sabouni I. Garrido D. Ramos J. M. 2008Crop losses due to olive moth mediated by ethylene. Pest Management Science 64 720 724 1526-4998
  113. 113. Reay-Jones F. P. F. Way M. O. Setamou M. Legendre B. L. Reagan T. E. 2003Resistance to the Mexican rice borer (Lepidoptera: Crambidae) among Louisiana and Texas sugarcane cultivars. Journal of Economic Entomology 96 1929 1934 0022-0493
  114. 114. Reay-Jones F. P. F. Showler A. T. Reagan T. E. Legendre B. L. Way M. O. Moser E. B. 2005Integrated tactics for managing the Mexican rice borer (Lepidopotera: Crambidae) in sugarcane. Environmental Entomology 34 1558 1565 0004-6225X..
  115. 115. Reay-Jones F. P. F. Wilson L. T. Showler A. T. Reagan T. E. Way M. O. 2007Role of oviposition on preference in an invasive crambid impacting two graminaceous host crops. Environmental Entomology 36 938 951 0004-6225X.
  116. 116. Reay-Jones F. P. F. Wilson L. T. Reagan T. E. Legendre B. L. Way M. O. 2008Predicting economic losses from the continued spread of the Mexican rice borer (Lepidoptera: Crambidae). Journal of Economic Entomology 101 237 250 0022-0493
  117. 117. Renwicke J. A. A. Radke C. D. 1988Sensory cues in host selection for oviposition by the cabbage butterfly, Pieris rapae. Journal of Insect Physiology 34 251 257 0022-1910
  118. 118. Renwicke J. A. A. Chew F. S. 1994Oviposition behavior in Lepidoptera. Annual Review of Entomology 39 377 400 0066-4170
  119. 119. Salama H. S. Rizk A. F. Sharaby A. 1984Chemical stimuli in flowers and leaves of cotton that affect behavior in the cotton moth, Spodoptera littoralis (Lepidoptera: Noctuidae). Entomologia 10 27 34
  120. 120. Schoonhoven L. M. 1981Chemical mediators between plants and phytophagous insects. In Semiochemicals: Their Role in Pest Control, Nordland, D.A.; Jones, R.L. & Lewis, W.L. (eds.), 31 50John Wiley & Sons, 100471058033York, New York, USA.
  121. 121. Schubert S. Serraj R. Plies-Balzer E. Mengel K. 1995Effect of drought stress on growth, sugar concentrations and amino acid accumulations in N 2 -fixing alfalfa (Medicago sativa). Plant Physiology 146 541 546 0981-9494
  122. 122. Schur K. Holdaway F. G. 1970Olfactory responses of female Ostrinia nubilalis (Lepidoptera: Pyraustinae). Entomologia Experimentalis et Applicata 13 455 461 0013-8703
  123. 123. Scriber J. M. 1977Limiting effects of low leaf-water content on the nitrogen utilization, energy budget, and larval growth of Hylophora cecropia. Oecologia 28 269 287 0029-8549
  124. 124. Seki M. Narusaka M. Abe H. Ksuga K. Yamaguchi-Shinozaki K. Carninci P. Hayashizaki P. Shinozaki K. 2001Monitoring the expression pattern of 1300 Arabidopsis genes under drought and cold stress by using a full-length cDNA microarray. Plant Cell Environment 13 61 72 1365-3040
  125. 125. Sharpe P. J. H. Wu H. I. Cates R. G. Goeschl J. D. 1985Energetics of pine defense systems to bark beetle attack., 206 223In Integrated Pest Management Research Symposium: the Proceedings, Branham, S.J. & Thatcher, R.C. (eds.), pp. 206-223. United States Department of Agriculture Forest Service General Technical Report SO-S6, Washington, DC, USA.
  126. 126. Showler A. T. Knaus R. M. Reagan T. E. 1989Foraging territoriality of the imported fire ant, Solenopsis invicta Buren, in sugarcane as determined by neutron activation analysis. Insectes Sociaux 36 235 239 0020-1812
  127. 127. Showler A. T. Reagan T. E. 1991Effects of sugarcane borer, weed, and nematode cottrol strategies in Louisiana sugarcane. Environmental Entomology 20 358 370 0004-6225X.
  128. 128. Showler A. T. Greenberg S. M. 2003Effects of weeds on selected arthropod herbivore and natural enemy populations, and on cotton growth and yield. Environmental Entomology 32 39 50 0004-6225X.
  129. 129. Showler A. T. Moran P. J. 2003Effects of drought stressed cotton, Gossypium hirsutum L., on beet armyworm, Spodoptera exigua (Hübner), oviposition, and larval feeding preferences and growth. Journal of Chemical Ecology 95 1971 1985 0098-0331
  130. 130. Showler A. T. Cavazos C. O. Moran P. J. 2007Dynamics of free amino acid accumulations in cotton leaves measured on different timelines after irrigation. Subtropical Plant Science 59 38 55 1009-7791
  131. 131. Showler A. T. Castro B. A. 2010aInfluence of drought stress on Mexican rice borer (Lepidoptera: Crambidae) oviposition preference in sugarcane. Crop Protection 28 722 727 0261-2194
  132. 132. Showler A. T. Castro B. A. 2010bMexican rice borer (Lepidoptera: Crambidae) oviposition site selection stimuli on sugarcane, and potential field applications. Journal of Economic Entomology 103 1180 1186 0022-0493
  133. 133. Showler A. T. Beuzelin J. M. Reagan T. E. 2011Alternate crop and weed host plant oviposition preferences by the Mexican rice borer (Lepidoptera: Crambidae). Crop Protection 30 895 901 0261-2194
  134. 134. Showler A. T. 2012Drought and arthropod pests of crops. In Drought: New Research, Neves, D.F & Sanz, J.D. (eds.), 131 154Nova Science, 978-1-62100-769-2Hauppauge, New York, USA.
  135. 135. Showler A. T. Reagan T. E. 2012Ecology and tactics for control of three sugarcane stalkboring species in the Western Hemisphere and Africa. In Sugarcane: Production, Cultivation and Uses, Goncalves, J.F. & Correia, K.D. (eds.), 1 32Nova Science, 978-1-61942-213-1Hauppauge, New York, USA.
  136. 136. Sidhu H. S. Kaur P. 1976The influence of water stress in the host plant on the reproduction of the mustard aphid, Lipaphis erysimi (Kalt.) Indian Journal of Ecology 3 163 166 0304-5250
  137. 137. Smitley D. R. Peterson N. C. 1996Interactions of water stress, honeylocust spider mites (Acari: Tetranychidae), early leaf abscission, and growth of Gleditsia tnacanthos. Journal of Economic Entomology 89 1577 1581 0022-0493
  138. 138. Städler E. 1984Contact chemoreception. In Chemical Ecology of Insects, Bell, W.J. & Carde, R.T. (eds.), 3 35Sinauer, Sunderland, Massachusetts, USA.
  139. 139. Stotz H. U. Pittendrigh B. R. Kroyman J. Weniger K. Fritsche J. Bauke A. Mitchell-Olds T. 2000Induced plant defense responses against chewing insects. Ethylene signaling reduces resistance of Arabidopsis against Egyptian cotton worm but not diamondback moth. Plant Physiology 124 1007 1018 0981-9428
  140. 140. Sturm M. M. Sterling W. L. 1990Geographical patterns of boll weevil mortality: observations and hypotheses. Environmental Entomology 19 59 65 0004-6225X.
  141. 141. Sumner L. C. Need J. T. Mc New R. W. Dorschner K. W. Elkenbary R. D. Johnson R. C. 1983Response of Schizaphis graminum (Homoptera: Aphididae) to drought-stressed wheat, using polyethylene glycol as ammatricum. Environmental Entomology 12 919 922 0004-6225X.
  142. 142. Udayagiri S. Mason C. E. 1995Host plant constituents as oviposition stimulants for a generalist herbivore: European corn borer. Entomologia Experimentalis et Applicata 76 59 65 0013-8703
  143. 143. Van Leerdam M. B. Johnson K. J. R. Smith J. W. Jr 1984Effects of substrate physical characteristics and orientation on oviposition by Eoreuma loftini (Lepidoptera: Pyralidae). Environmental Entomology 13 800 802 0004-6225X.
  144. 144. Van Leerdam M. B. Johnson K. J. R. Smith J. W. Jr 1986Ovipositional sites of Eoreuma loftini (Lepidoptera: Pyralidae) in sugarcane. Environmental Entomology 15 75 78 0004-6225X.
  145. 145. Van Zwaluenberg R. H. 1926Insect enemies of sugarcane in western Mexico. Journal of Economic Entomology 19 664 669 0022-0493
  146. 146. Vanderzant E. S. 1958The amino acid requirements of the pink bollworm. Journal of Economic Entomology 51 309 311 0022-0493
  147. 147. Vincent D. Lapierre C. Pollet B. Cornic G. Negroni L. Zivy M. 2005Water deficits affect caffeate O-methyltransferase, lignifications, and related enzymes in maize leaves. A proteomic investigation. Plant Physiology 137 949 960
  148. 148. Vité J. P. Volz H. A. Paiva M. R. Bakke A. 1986Semiochemicals in host selection and colonization of pine trees by the pine shoot beetle Tomicus piniperda. Naturwissenschaften 73 39 40 0028-1042
  149. 149. Waladde S. M. 1983Chemoreceptors of adult stem borers: tarsal and ovipositor sensilla on Chilo partellus and Eldana saccharina. Insect Science and its Applications 4 159 165 0191-9040
  150. 150. Wearing C. H. Van Emden H. F. 1967Studies on the relations of insect and host plant. I. Effects of water stress in host plants on infestations by Aphis fabae Scop., Myzus persicae (Sulz.) and Brevicoryne brassicae (L.). Nature 213 1051 1052 0028-0836
  151. 151. Weibull J. 1987Seasonal changes in the free amino acids of oat and barley phloem sap in relation to plant growth stage and growth of Rhopalosiphum padi. Annals of Applied Biology 111 727 737 1744-7348
  152. 152. White T. C. R. 1984The abundance of invertebrate herbivores in relation to the availability of nitrogen in stressed food plants. Oecologia 63 90 105 0029-8549
  153. 153. Wilson B. 2011Advanced management of the Mexican rice borer (Eoreuma loftini) in sugarcane. M.S. thesis, Louisiana State University, Baton Rouge, Louisiana, USA.
  154. 154. Wright L. C. Berryman A. A. Gurusiddaiah S. 1979Host resistance to the fir engraver beetle, Scolytus ventralis (Coleoptera: Scolytidae). IV. Effe cts of defoliation on wound monoterpene and inner bark carbohydrate concentrations. Canadian Entomologist 111 1255 1262 0000-8347X.
  155. 155. Zalucki M. P. Clarke A. R. Malcom S. B. 2002Ecology and behavior of first instar larval Lepidoptera. Annual Review of Entomology 47 361 393 0066-4170
  156. 156. Zhang J. X. Kirkham M. B. 1990Variation in ethylene production by sorghum. Euphytica 46 109 117 0014-2336
  157. 157. Zhu J. K. 2002Salt and drought stress signal transduction in plants. Annual Review of Plant Biology 53 247 273 1543-5008

Written By

Allan T. Showler

Submitted: 01 May 2012 Published: 13 March 2013