\r\n\tThis book will present and discuss the advancement of research on age-associated diseases and their underlying mechanisms, exploring mainly causal relation aspects of the glutathione peroxidase.
",isbn:"978-1-83880-126-7",printIsbn:"978-1-83880-125-0",pdfIsbn:null,doi:null,price:0,priceEur:0,priceUsd:0,slug:null,numberOfPages:0,isOpenForSubmission:!1,hash:"127defed0a50ad5ed92338dc96e1e10e",bookSignature:"Dr. Margarete Bagatini",publishedDate:null,coverURL:"https://cdn.intechopen.com/books/images_new/9002.jpg",keywords:"Free Radicals, Antioxidants, Health, Glutathione Peroxidase, Superoxide Dismutase, Catalase, Structure, Activity, Infectious Diseases, Coronary Diseases, Neurological Diseases, Protection",numberOfDownloads:54,numberOfWosCitations:0,numberOfCrossrefCitations:0,numberOfDimensionsCitations:0,numberOfTotalCitations:0,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"August 27th 2019",dateEndSecondStepPublish:"September 17th 2019",dateEndThirdStepPublish:"November 16th 2019",dateEndFourthStepPublish:"February 4th 2020",dateEndFifthStepPublish:"April 4th 2020",remainingDaysToSecondStep:"3 months",secondStepPassed:!0,currentStepOfPublishingProcess:4,editedByType:null,kuFlag:!1,editors:[{id:"217850",title:"Dr.",name:"Margarete",middleName:null,surname:"Bagatini",slug:"margarete-bagatini",fullName:"Margarete Bagatini",profilePictureURL:"https://mts.intechopen.com/storage/users/217850/images/system/217850.jpeg",biography:"Margarete Dulce Bagatini has been an Associate Professor at the Federal University of Fronteira Sul (UFFS) since 2011. She has graduated from Federal University of Santa Maria / UFSM (Pharmacy - Clinical Analysis, 2006) and she obtained Ph.D. (2010) in Biological Sciences from the same university. She is a member of the research advisory committee (2013), assistant coordinator of research (2014) and academic coordinator (2015) of Campus Chapecó /UFFS. Also she is a leader of the research group: Biological and Clinical Studies in Human Pathologies, professor of postgraduate programs and member of the Committee UFFS Research Advisor. She has experience in the field of Pharmacy, Clinical Analysis, Microbiology, Immunology and Biochemistry, working mainly in the following subjects: oxidative stress, purinergic system and human pathologies. She has been a reviewer of several international journals and editor of the journals Journal of Immunology Research and Oxidative Medicine and Cellular Longevity.",institutionString:"Universidade Federal da Fronteira Sul",position:null,outsideEditionCount:0,totalCites:0,totalAuthoredChapters:"1",totalChapterViews:"0",totalEditedBooks:"0",institution:{name:"Universidade Federal da Fronteira Sul",institutionURL:null,country:{name:"Brazil"}}}],coeditorOne:null,coeditorTwo:null,coeditorThree:null,coeditorFour:null,coeditorFive:null,topics:[{id:"6",title:"Biochemistry, Genetics and Molecular Biology",slug:"biochemistry-genetics-and-molecular-biology"}],chapters:[{id:"69266",title:"Subcellular Localization of Glutathione Peroxidase, Change in Glutathione System during Ageing and Effects on Cardiometabolic Risks and Associated Diseases",slug:"subcellular-localization-of-glutathione-peroxidase-change-in-glutathione-system-during-ageing-and-ef",totalDownloads:54,totalCrossrefCites:0,authors:[null]}],productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"},personalPublishingAssistant:{id:"220812",firstName:"Lada",lastName:"Bozic",middleName:null,title:"Ms.",imageUrl:"https://mts.intechopen.com/storage/users/220812/images/6021_n.jpg",email:"lada@intechopen.com",biography:"As an Author Service Manager my responsibilities include monitoring and facilitating all publishing activities for authors and editors. 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\n
1. Introduction
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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\n\n 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; \n Showler, 2012\n ) 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.
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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.
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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
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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; \n Showler, 2012\n\n ). 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.
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; \n Showler, 2012\n\n ). 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, \n 2012\n\n ; 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).
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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.
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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
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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 (\n Showler, 2012\n ). Black bean aphid, Aphis\n\n 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\n 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).
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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; \n Showler & Moran, 2003\n\n ). 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
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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; \n Showler, 2012\n\n ), 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 (\n Showler, 2012\n\n ), 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
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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 (\n Johnson, 1981\n , 1984; \n Johnson & Van Leerdam, 1981\n\n ), 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 (\n Showler & Reagan, 2012\n ).
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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).
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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).
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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).
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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).
\n
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 (\n Johnson, 1981\n\n ); 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).
\n
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.
\n
\n
\n
\n
\n \n
\n
\n
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)
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).
\n a Cystine was detectable but not found in the samples.
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 (\n Showler, 2012\n ; \n Showler & Reagan, 2012\n\n ). 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 (\n Showler & Greenberg, 2003\n\n ). 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 (\n Showler & Reagan, 2012\n ) despite its cryptic larval behavior.
\n
\n
\n
6. Conclusion
\n
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.
\n
\n \n',keywords:null,chapterPDFUrl:"https://cdn.intechopen.com/pdfs/43230.pdf",chapterXML:"https://mts.intechopen.com/source/xml/43230.xml",downloadPdfUrl:"/chapter/pdf-download/43230",previewPdfUrl:"/chapter/pdf-preview/43230",totalDownloads:2217,totalViews:336,totalCrossrefCites:0,totalDimensionsCites:11,hasAltmetrics:0,dateSubmitted:"May 1st 2012",dateReviewed:"September 6th 2012",datePrePublished:null,datePublished:"March 13th 2013",readingETA:"0",abstract:null,reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/43230",risUrl:"/chapter/ris/43230",book:{slug:"abiotic-stress-plant-responses-and-applications-in-agriculture"},signatures:"Allan T. Showler",authors:[{id:"72273",title:"Dr.",name:"Allan T.",middleName:null,surname:"Showler",fullName:"Allan T. Showler",slug:"allan-t.-showler",email:"allan.showler@ars.usda.gov",position:null,institution:{name:"Agricultural Research Service",institutionURL:null,country:{name:"United States of America"}}}],sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_2",title:"2. Water deficit, host plant nutrient accumulation, and associations with phytophagous arthropods",level:"1"},{id:"sec_3",title:"3. Water is a nutrient, too",level:"1"},{id:"sec_4",title:"4. Some non-nutrient-related associations of water deficit with phytophagous arthropods",level:"1"},{id:"sec_5",title:"5. Multiple effects of water deficit: case study on sugarcane and the Mexican rice borer",level:"1"},{id:"sec_6",title:"6. Conclusion",level:"1"}],chapterReferences:[{id:"B1",body:'\n \n \n \n Akbar\n W.\n \n \n Showler\n A. T.\n \n \n White\n W. H.\n \n \n Reagan\n T. E.\n \n \n 2010Categorizing sugarcane cultivars for resistance to the sugarcane aphid and yellow sugarcane aphid (Hemiptera: Aphididae). Journal of Economic Entomology\n 103\n 1431\n 1437\n 0022-0493\n \n '},{id:"B2",body:'\n \n \n \n Apelbaum\n A.\n \n \n Tang\n S. 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M.\n \n \n 2003Effects of weeds on selected arthropod herbivore and natural enemy populations, and on cotton growth and yield. Environmental Entomology 32\n 39\n 50\n 0004-6225X.\n '},{id:"B129",body:'\n \n \n \n Showler\n A. T.\n \n \n Moran\n P. J.\n \n \n 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\n 1971\n 1985\n 0098-0331\n \n '},{id:"B130",body:'\n \n \n \n Showler\n A. T.\n \n \n Cavazos\n C. O.\n \n \n Moran\n P. J.\n \n \n 2007Dynamics of free amino acid accumulations in cotton leaves measured on different timelines after irrigation. Subtropical Plant Science 59\n 38\n 55\n 1009-7791\n \n '},{id:"B131",body:'\n \n \n \n Showler\n A. T.\n \n \n Castro\n B. A.\n \n \n 2010aInfluence of drought stress on Mexican rice borer (Lepidoptera: Crambidae) oviposition preference in sugarcane. Crop Protection 28\n 722\n 727\n 0261-2194\n \n '},{id:"B132",body:'\n \n \n \n Showler\n A. T.\n \n \n Castro\n B. A.\n \n \n 2010bMexican rice borer (Lepidoptera: Crambidae) oviposition site selection stimuli on sugarcane, and potential field applications. Journal of Economic Entomology\n 103\n 1180\n 1186\n 0022-0493\n \n '},{id:"B133",body:'\n \n \n \n Showler\n A. T.\n \n \n Beuzelin\n J. M.\n \n \n Reagan\n T. E.\n \n \n 2011Alternate crop and weed host plant oviposition preferences by the Mexican rice borer (Lepidoptera: Crambidae). Crop Protection\n 30\n 895\n 901\n 0261-2194\n \n '},{id:"B134",body:'\n \n \n \n Showler\n A. T.\n \n \n 2012Drought and arthropod pests of crops. In Drought: New Research, Neves, D.F & Sanz, J.D. (eds.), 131\n 154Nova Science, 978-1-62100-769-2Hauppauge, New York, USA.\n '},{id:"B135",body:'\n \n \n \n Showler\n A. T.\n \n \n Reagan\n T. E.\n \n \n 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\n 32Nova Science, 978-1-61942-213-1Hauppauge, New York, USA.\n '},{id:"B136",body:'\n \n \n \n Sidhu\n H. S.\n \n \n Kaur\n P.\n \n \n 1976The influence of water stress in the host plant on the reproduction of the mustard aphid, Lipaphis erysimi (Kalt.) Indian Journal of Ecology\n 3\n 163\n 166\n 0304-5250\n \n '},{id:"B137",body:'\n \n \n \n Smitley\n D. R.\n \n \n Peterson\n N. C.\n \n \n 1996Interactions of water stress, honeylocust spider mites (Acari: Tetranychidae), early leaf abscission, and growth of Gleditsia tnacanthos. Journal of Economic Entomology\n 89\n 1577\n 1581\n 0022-0493\n \n '},{id:"B138",body:'\n \n \n \n Städler\n E.\n \n \n 1984Contact chemoreception. In Chemical Ecology of Insects, Bell, W.J. & Carde, R.T. (eds.), 3\n 35Sinauer, Sunderland, Massachusetts, USA.\n '},{id:"B139",body:'\n \n \n \n Stotz\n H. U.\n \n \n Pittendrigh\n B. R.\n \n \n Kroyman\n J.\n \n \n Weniger\n K.\n \n \n Fritsche\n J.\n \n \n Bauke\n A.\n \n \n Mitchell-Olds\n T.\n \n \n 2000Induced plant defense responses against chewing insects. Ethylene signaling reduces resistance of Arabidopsis against Egyptian cotton worm but not diamondback moth. Plant Physiology 124\n 1007\n 1018\n 0981-9428\n \n '},{id:"B140",body:'\n \n \n \n Sturm\n M. M.\n \n \n Sterling\n W. L.\n \n \n 1990Geographical patterns of boll weevil mortality: observations and hypotheses. Environmental Entomology\n 19\n 59\n 65\n 0004-6225X.\n '},{id:"B141",body:'\n \n \n \n Sumner\n L. C.\n \n \n Need\n J. T.\n \n \n Mc New\n R. W.\n \n \n Dorschner\n K. W.\n \n \n Elkenbary\n R. D.\n \n \n Johnson\n R. C.\n \n \n 1983Response of Schizaphis graminum (Homoptera: Aphididae) to drought-stressed wheat, using polyethylene glycol as ammatricum. Environmental Entomology\n 12\n 919\n 922\n 0004-6225X.\n '},{id:"B142",body:'\n \n \n \n Udayagiri\n S.\n \n \n Mason\n C. E.\n \n \n 1995Host plant constituents as oviposition stimulants for a generalist herbivore: European corn borer. Entomologia Experimentalis et Applicata\n 76\n 59\n 65\n 0013-8703\n \n '},{id:"B143",body:'\n \n \n \n Van Leerdam\n M. B.\n \n \n Johnson\n K. J. R.\n \n \n Smith\n J. W.\n \n \n Jr \n \n \n \n 1984Effects of substrate physical characteristics and orientation on oviposition by Eoreuma loftini (Lepidoptera: Pyralidae). Environmental Entomology\n 13\n 800\n 802\n 0004-6225X.\n '},{id:"B144",body:'\n \n \n \n Van Leerdam\n M. B.\n \n \n Johnson\n K. J. R.\n \n \n Smith\n J. W.\n \n \n Jr \n \n \n \n 1986Ovipositional sites of Eoreuma loftini (Lepidoptera: Pyralidae) in sugarcane. Environmental Entomology 15\n \n 75\n 78\n 0004-6225X.\n '},{id:"B145",body:'\n \n \n \n Van Zwaluenberg\n R. H.\n \n \n 1926Insect enemies of sugarcane in western Mexico. Journal of Economic Entomology\n 19\n 664\n 669\n 0022-0493\n \n '},{id:"B146",body:'\n \n \n \n Vanderzant\n E. S.\n \n \n 1958The amino acid requirements of the pink bollworm. Journal of Economic Entomology\n 51\n 309\n 311\n 0022-0493\n \n '},{id:"B147",body:'\n \n \n \n Vincent\n D.\n \n \n Lapierre\n C.\n \n \n Pollet\n B.\n \n \n Cornic\n G.\n \n \n Negroni\n L.\n \n \n Zivy\n M.\n \n \n 2005Water deficits affect caffeate O-methyltransferase, lignifications, and related enzymes in maize leaves. A proteomic investigation. Plant Physiology\n 137\n 949\n 960\n \n '},{id:"B148",body:'\n \n \n \n Vité\n J. P.\n \n \n Volz\n H. A.\n \n \n Paiva\n M. R.\n \n \n Bakke\n A.\n \n \n 1986Semiochemicals in host selection and colonization of pine trees by the pine shoot beetle Tomicus piniperda. Naturwissenschaften\n 73\n 39\n 40\n 0028-1042\n \n '},{id:"B149",body:'\n \n \n \n Waladde\n S. M.\n \n \n 1983Chemoreceptors of adult stem borers: tarsal and ovipositor sensilla on Chilo partellus and Eldana saccharina. Insect Science and its Applications\n 4\n 159\n 165\n 0191-9040\n \n '},{id:"B150",body:'\n \n \n \n Wearing\n C. H.\n \n \n Van Emden\n H. F.\n \n \n 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\n 213\n 1051\n 1052\n 0028-0836\n \n '},{id:"B151",body:'\n \n \n \n Weibull\n J.\n \n \n 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\n 727\n 737\n 1744-7348\n \n '},{id:"B152",body:'\n \n \n \n White\n T. C. R.\n \n \n 1984The abundance of invertebrate herbivores in relation to the availability of nitrogen in stressed food plants. Oecologia 63\n 90\n 105\n 0029-8549\n \n '},{id:"B153",body:'\n \n \n \n Wilson\n B.\n \n \n 2011Advanced management of the Mexican rice borer (Eoreuma loftini) in sugarcane. M.S. thesis, Louisiana State University, Baton Rouge, Louisiana, USA.\n '},{id:"B154",body:'\n \n \n \n Wright\n L. C.\n \n \n Berryman\n A. A.\n \n \n Gurusiddaiah\n S.\n \n \n 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\n 111\n 1255\n 1262\n 0000-8347X.\n '},{id:"B155",body:'\n \n \n \n Zalucki\n M. P.\n \n \n Clarke\n A. R.\n \n \n Malcom\n S. B.\n \n \n 2002Ecology and behavior of first instar larval Lepidoptera. Annual Review of Entomology 47\n 361\n 393\n 0066-4170\n \n '},{id:"B156",body:'\n \n \n \n Zhang\n J. X.\n \n \n Kirkham\n M. B.\n \n \n 1990Variation in ethylene production by sorghum. Euphytica 46\n \n 109\n 117\n 0014-2336\n \n '},{id:"B157",body:'\n \n \n \n Zhu\n J.\n \n \n \n K.\n \n \n 2002Salt and drought stress signal transduction in plants. Annual Review of Plant Biology\n 53\n 247\n 273\n 1543-5008\n \n '}],footnotes:[],contributors:[{corresp:null,contributorFullName:"Allan T. Showler",address:"Allan.Showler@ars.usda.gov",affiliation:'
USDA-ARS, Weslaco, Texas, USA
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1. Introduction
Teeth are a topic of interest to paleontologists because they are very well preserved. As a matter of fact, the dental remains have made it possible to study the evolution of mammals by analyzing their morphology. In developmental biology, the mouse model is an interesting model for studying dental development.
Humans have two dentitions (temporary and permanent) and different types of teeth, incisor, canine, premolar, and molar with different morphologies, whereas mice only have two types (incisor and molar) separated by a diastema from which the incisors have unlimited growth. Despite these differences, the dental development process is similar in humans and mice, and regulatory phenomena have been maintained over the evolution.
Teeth, such as mammary glands, hair, and feathers, develop from two adjacent tissues: the epithelium and the mesenchyme, although they all have different morphologies. Indeed, during development, the specific shape of each organ is defined in relation to epithelial-mesenchymal proliferation and to all the changes that the epithelium undergoes [1].
The embryological aspect of the molars was addressed in order to clarify the etiopathogenic aspect and to adapt therapeutic attitudes according to the diagnosis.
The objective of this chapter is to address the embryology of human molars by focusing on its molecular and morphological characteristics.
2. Phylogenetic aspects
Teeth represent a new morphological feature of mammals [2, 3]. Molars are complex teeth able to become occluded. Interlocking intercuspation between upper and lower molars allows food to be crushed [4]. Evolutionary dietary radiations are related to the great diversity of the current mammalian molars. They are clarified in the fossil record, where new molar organizations are often related to significant line diversifications. Several theories have been advanced to explain the evolution of molars. Like all primates, Man is a placental mammal, and the ancestor of contemporary humans is Homo sapiens. For 200 million years, in Therian mammals, the molars have trigonodontal morphology; in other words, the three tubercles are arranged in a triangle [5].
In 1965, the discovery of a fossil of a lower molar made it possible to show that on this Therian branch around 135 million years ago, these molars already existed. They were called tribosphenic by Simpson in 1936 [6]. These mandibular molars have six tubercles, three of which are pointed, high, sharp, and are arranged in a triangle and distal position. The three others tubercles are lower and are arranged in a central basin to receive the main palatal tubercle of the opposite teeth that have only three cusps. The fact of having six tubercles is of physiological interest when taking food.
Nearly 110 million years ago, the oldest placental mammals had a dental formula with 52 teeth, including 3 molars in a decreasing series, the first being the largest. This primitive disposition is found in modern man.
Around 75 million years ago, with the dinosaurs extinction, other species invaded space, and the dental formula was reduced to 44 teeth for all placental mammals including the man.
In the Catarrhini, the loss of one incisor and two premolars leads to a dental formula with 32 teeth found in monkeys of the ancient world (Afro-Eurasia), the Hominids, and the contemporary Men. It has been recognized for 45 million years [7].
In the genus Homo, the 32-teeth morphology does not differ much from the modern men, except for the great variability in size. Root morphology may vary from one group to another. The reduction in the number of cusps observed in humans can be considered as a specialization trait and not as a step backward. However, the reduction in the dental formula in the placentals and primates mainly affected the incisors, premolars, and even canines but not the molars.
Wisdom tooth agenesis, especially mandibular agenesis, is often considered as a sign of evolution. On the other hand, the presence of supernumerary teeth or hypergenesis is explained as a return to ancestral forms
3. Morphological aspects
3.1 Formation of the odontogenic epithelium
The odontogenic epithelium is formed from the oral epithelium that lines the primary oral cavity called the “stomodeum.” It appears as a localized thickening of the oral epithelium, and it is formed by several cellular layers resulting from a series of localized mitoses affecting the oral epithelium. The mitotic spindle of dividing cells is oriented perpendicular to the basal membrane that separates the epithelium from the ectomesenchyma.
3.2 Placement of the vestibular and primary dental blades
Epithelial thickening continues to proliferate and sinks into the underlying ectomesenchymal tissue forming a plunging wall (also called a primitive dental blade). This latter splits into two blades: vestibular and dental. The vestibular blade determines the formation of the buccal vestibule, which is the space between the cheek/lip and the dental arch.
3.3 Evolution of dental placodes
In humans, as in rats and mice, the dental blade will give birth to the dental placodes that will be at the origin of the formation of future dental germs. Dental placodes are cellular clusters attached to the dental blade by a net of epithelial cells called the primary dental blade. Each dental arch initially contains 10 dental placodes. From the primary dental blade develops the secondary dental blade, which is at the origin of the 16 permanent teeth per arch.
Each placode will undergo morphological changes that are described as three successive stages: bud stage, cup stage, and bell stage [1].
4. Placement of molar dental germs
Since the three molars are not preceded by temporary teeth, they evolve from the distal end of the initial dental blade, which proliferates in a posterior direction. The primary dental blade of the second temporary molar will cause the formation of four secondary dental blades. For each half of the arch, starting from the anterior area toward the posterior area, each of these four secondary dental blades will give the permanent germ of the following teeth: the first permanent molar, the second permanent molar, and the third permanent molar.
The secondary dental blades that are at the origin of the formation of the 1st and 2nd molar will orient themselves vertically as long as they have space that allows them to orient themselves in the mesenchyma. On the other hand, in most cases for the 3rd molar, orientation problems arise because there is not enough space for its secondary dental blade to be parallel to the other two blades [8].
All dental buds, with the exception of the second and third permanent molars, are present and begin to develop before birth [9]. The chronology of the appearance of molar germs remains variable according to the authors; however, it is often found that the germ of the first molar appears around the 4th or 5th month of intrauterine life. The one of the second molar appears around the 9th month or 1 year after birth.
The germ of the third molar does not appear until around 4 or 5 years of age. Mineralization begins between 7, 9, and 10 years, and the crown is completed between 12 and 16 years. The emergence in the oral cavity is between 17- and 21-year-olds; the tooth will then slide along the distal surface of the second molar to reach the occlusion level. Root building ends between the ages of 18 and 25 years. The place it has depends on the growth in the posterior region of the arch. The main activity of the dental blade is spread over a period of about 5 years. However, the dental blade near the third molar continues to be active until about 15 years of age [9].
A number of anomalies can occur during the development of the tooth. The development of excess dental blade can lead to an increase in the number of dental buds, resulting in too many teeth (supernumerary). A deficient dental blade can lead to a reduction in the number of teeth (hypodontia) [9].
5. Root formation
Molars are multiradiculated teeth. Indeed, the vast majority of the first maxillary molars have three roots. The second maxillary molar has more frequent variations in the number of roots than the first maxillary molar, and the first mandibular molar and the second have two roots in the majority.
Root formation or radiculogenesis or rhizagenesis is the development of the root pulpo-dentinary organ in close relationship with cemenesis, the outline of the dentoalveolar ligament and the construction of the alveolar bone. It begins when the final dimensions are acquired. The Hertwig epithelial sheath is at the origin of root formation, depending on their number, shape, and size [10].
As for the crown, root development is governed by interactions involving the Hertwig epithelial sheath, basement membrane, mesenchymal papilla, and dental follicle.
5.1 Formation of the Hertwig epithelial sheath
The Hertwig epithelial sheath originates from the reflection zone or cervical loop which is the place where the external and internal adamantin epitheliums (EAE and EAI) meet to form a double epithelial layer. Hertwig epithelial sheath has an annular structure surrounded by a basal membrane that separates it from the pulpal and follicular mesenchyma. This basement membrane has anchoring fibrils on the pulp side. The internal epithelium faces the papilla and the external epithelium faces the dental follicle. The Hertwig epithelial sheath will emit tongues in the centripetal direction that will fuse in the central region of the papilla and form rings from which the roots can be identified. The number of strips emitted is proportional to the number of roots that each molar can have. For example, for the molar which will have two roots, two tongues are formed, and after fusion of two rings, each of the two will be at the origin of the formation of a root. These two leaves remain attached and progress in the underlying connective tissue in the apical direction defining the future shape of the dental root [11].
Root elongation and tissue formation are related to the coordinated proliferation of sheath epithelial cells and surrounding mesenchymal cells [12].
5.2 Formation of root dentin, cement, and apex
Root dentin forms in parallel with the proliferation in the apical direction of the Hertwig sheath. The latter gradually induces odontoblastic differentiation. The pulp parenchyma cells close to the anchor fibrils differentiate into odontoblasts. These odontoblasts produce preentine, which mineralizes to form dentin. The cells of the outer dental epithelium forming the outer layer of the sheath do not differentiate into ameloblasts as is the case for the crown. Then, the basement membrane degrades, and the epithelial blade involutes and gradually dissociates.
Developmental defects of the Hertwig sheath at the apical third of the root are at the origin of the formation of the lateral canals following a stop of dentinogenesis at this site due to the nondifferentiation of pulp fibroblasts into odontoblasts.
The cells of the sheath can undergo three spells: some can form the “Malassez epithelial debris,” others can die by apoptosis, while others can undergo epithelial-mesenchymal transformation.
As the sheath disintegrates, follicular cells near the surface of the root dentin differentiate into cementoblasts. These synthesize and deposit the cement matrix in contact with the dentin.
As the root development progresses, the epithelial ring forming the Hertwig epithelial sheath gradually shrinks as a result of a reduction in mitosis, thereby reducing the size of the root tube. This narrowing allows the development of one or more orifices (or foramina), which are the place where vascular and nervous elements intended for the pulp to pass through.
The development of the root ends with the construction of the apex, which is a slow process. In humans, for example, for the 1st permanent molar, this operation is performed until the age of 9–10 years. In the case of permanent teeth, this phenomenon lasts longer and requires more time than the development of the root itself.
6. Molecular aspects
6.1 Epithelial-mesenchymal interactions
In humans, dental development includes the morphogenesis of crowns and roots and results in the formation of the enamel organ, odontoblastic, ameloblastic, and cementoblastic differentiation. Huge advances in research have made it possible to understand the phenomena of molecular regulation of dental development.
Dental development follows a precisely controlled and regulated genetic program. The dental organ consists of an epithelial part that derives from the ectoderm and a mesenchymal part that derives from mesodermal cells on the one hand and cells from neural ridges on the other hand [13, 14, 15, 16].
The dental organ develops from a communication between the epithelium and the underlying mesenchyma. The communication language has been preserved throughout the evolution. This communication between the epithelium and the mesenchyma is done through signaling molecules and growth factors [17, 18, 19].
The studies carried out on the mouse molar have enabled us to gather a body of knowledge with many similarities to those of humans. However, the experimental data obtained in animals can be extrapolated relatively reliably to understand what is actually happening in humans.
Several families have been described, including:
TGF-beta (transforming growth factor beta) including BMP (bone morphogenetic proteins) activins and follistatin;
FGF (Fibroblast growth factors);
Hedgehog (only Sonic hedgehog (Shh) is known for its role in odontogenesis);
These molecules send their message to the nucleus through the signaling pathways and receptors on the cell membrane surface. Transcription factors will then modulate the expression of different target genes and induce changes in cell response and behavior (Figure 1) [25].
Figure 1.
Signaling in tooth development [25].
Genes represented in “light blue” colored squares or rectangles are responsible, when inactivated, for stopping dental development.
6.2 Determination of the dental region
It should be remembered that the odontogenic epithelium is formed at the first gill arch. The latter undergoes pharyngeal regionalization, resulting in the expression of Fgf8 and 9 (fibroblast growth factors 8 and 9) and Lhx-6 and -7 (LIM homeobox 6 and 7) in the oral part (rostral) and Gsc (goosecoid) in the aboral part (caudal). Indeed, the expression of Fgf8 in the odontogenic epithelium in the oral part of the first pharyngeal arch causes the expression of Lhx-7 in the underlying ectomesenchyma. In the aboral region, there is an important expression of Gsc in the ectomesenchyma. Gsc expression in the caudal region is not responsible for inhibiting Lhx-7 expression in this area; however, Lhx-7 expression in the rostral region will result in blocking Gsc gene expression in this.
In addition to Fgf8, a second BMP4 signaling molecule (bone morphogenetic protein 4) is expressed in the epithelium in the distal and therefore in the median region of the 1st arc.
The activation and inhibition of transcription factors allows the delimitation of the odontogenic territory by BMP4 and Fgf8a double signalling. (Figure 2) [19].
Figure 2.
Pattern of gene expression in the developing tooth [19]. (a) Signaling within the epithelium and between the epithelium and the mesenchyme at embryonic day (E) 10.5. The diagram shows an isolated mandibular arch. Positive autoregulatory loops and mutual repression within the epithelium lead to the formation of strict boundaries of gene expression, which sets up the presumptive incisor and molar fields. Members of the bone morphogenetic protein (BMP) and fibroblast growth factor (FGF) families of protein in the epithelium induce and inhibit the expression of various homeobox genes. This results in a complex pattern of gene expression in the mesenchyme, across both the proximal–distal and oral–aboral/rostral–caudal axes. (b) The odontogenic homeobox code model of dental patterning. The nested expression pattern of homeobox genes in the mandible produces a homeobox code that defines tooth type. Bapx1 (bagpipe homeobox gene 1 homolog); Barx1 (BarH-like homeobox 1); Dlx (distalless homeobox); Gsc (goosecoid); Lhx (LIM homeodomain genes); Msx (homeobox, msh-like); Pitx (paired-related homeobox gene).
6.3 Determination of dental identity
Mammalian teeth are meristic series. The determination of different morphology was explained by two theories:
The gradient theory proposed by Butler [26] which stipulates the presence of morphogenetic fields and that the determination of the shape of the tooth is a function of its position in the field independent of local factors.
The theory of clones proposed by Osborn [27] which stipulates that ectomesenchyma is already differentiated into three cellular clones, incisal, canine, and molar clones, before its migration. The proposal of this second concept suggested that the two theories are competing.
In 1995, the theory of odontogenic homeocode was developed by Sharpe [22], which represents a synthesis of the two theories: gradients and clones and shows that the latter two are complementary. These two concepts were explained in the light of the discovery of new genes and signaling molecules (Figure 3) [26, 27, 28].
The identity of each tooth, including the molars, is characterized by its homeocode, which represents the combination of homeogens that defines the position and identity of the tooth. Indeed, different homeogens are expressed by the neural crest cells of the ectomesenchyma under the instructive induction of the oral epithelial cells. These homeogens are divergent and therefore of the nonhox type.
This odontogenic homeocode theory involves four homogenous genes: muscle segment homeodomain-homeobox 1 (Msx-1), muscle segment homeodomain-homeobox 2 (Msx-2), distal-less homeobox 1 (Dlx1), and goosecoide. In the molar sector, Msx-1 and Dlx-1 are expressed and Msx-2 and goosecoide are not expressed. In the canine sector, Msx-1, Msx-2, and goosecoide are expressed, and Dlx-1 is not expressed; in incisal sector, Msx-1 and goosecoide are expressed, Msx-2 and Dlx-1 are not expressed.
In the concept of morphogenetic fields, the consideration of various genetic factors and their epigenetic modulation influences dental development [29].
According to Mitsiadis’ work in 2006, the three models, gradients, clones, and homeocodes, could be grouped into a single model to explain dental identity. Indeed, dental identity, including molars, is given by the presence of morphogenetic fields defined by the diffusion of growth factors. The odontogenic epithelium expresses gradients of signaling molecules that are mainly Fgf, Bmp, Shh, and Wint that will diffuse to the underlying mesenchymal tissue containing neural peak cells. Depending on the location and instruction received by these cells, they will express a set of divergent genes in relation to concentrations of signaling molecules. The locally defined tooth type is related to the locally expressed divergent homeogen combinatorics of these ridge cells (Figure 4) [30].
Figure 4.
Dental identity determination (adapted from Ref. [30]).
The Mitsiadis model combines the three concepts: morphogenetic fields, clone, and odontogenic homeocode.
These three models should be viewed as complementary rather than contradictory and propose that this unifying view can be extended into the clinical setting using findings on dental patterning in individuals with missing teeth. The proposals are compatible with the unifying etiological model developed by Brook in 1984 based on human epidemiological and clinical findings. Indeed, this new synthesis can provide a sound foundation for clinical diagnosis, counseling, and management of patients with various anomalies of dental development, as well as suggesting hypotheses for future studies.
6.4 Molecular factors involved in root formation
The root development process involves a set of signaling cascades. Various growth factors, including BMPs (bone morphogenetic proteins), EGF (epidermal growth factor), IGF (insulin-like growth factor), FGF (fibroblast growth factor), transcription factors Msx1, Msx2, Runx-2, Sonic Hedgehog (Shh), enamel proteins (secreted by HGH cells), and other proteins such as follistatin and activin A, are involved in the root development process. Indeed, they are involved in the growth and/or differentiation of odontoblasts and cementoblasts and/or in the mineralization of dentin and/or cementum [21, 31, 32, 33, 34, 35, 36].
7. Signaling center (primary and secondary enamel knots)
Dental morphology is controlled by an epithelial signaling center called the enamel node. The node of the enamel is a particular and transient histological structure formed by a cellular cluster that appears at the basal part of the internal dental epithelium. The node of the primary enamel is present in the dental germs of all types of teeth including incisors.
Because the enamel nodes link cell differentiation to morphogenesis, Thesleff suggests that the latter can be considered as central regulators of dental development [37].
During molar development, the node of the secondary enamel is formed during the bell stage at the location of future cusp areas. At this point, the expression of signaling molecules precedes the folding and growth of the dental epithelium [38, 39].
The Slit1 gene is expressed in the nodes of the primary and secondary enamel during the formation of molar cusps [40].
8. Genes and dental problems
The approaches provided by Line and Mitsiadis have advanced the clinic’s understanding of dental identity establishment based on gradient, clone, and homeocode theories [29, 30].
The multifactorial model involving genetic, epigenetic, and environmental determinants has provided better explanations and helped to understand missing and supernumerary teeth in monozygotic twins [41].
In humans, dental problems are observed during pathologies of dental development or syndromes.
Mutations in genes known as divergent homeobox genes encoding transcription factors such as MSX1 and PAX9 (paired domain box gene 9) are at the origin of oligodontia. Indeed, a mutation in the homeobox of the MSX1 gene (substitution of an arginine by a proline in the homeodomain region) is associated with the agenesis of third molars, indicating the involvement of MSX1 in the dentition pattern [42, 43, 44].
Also, mutations in the PAX9 gene cause oligodontia characteristic of molars [45, 46, 47, 48]. The severity of dental agenesis appears to be correlated with the ability of the mutated PAX9 protein to bind to DNA [49].
A misdirection mutation during the sequencing of the PAX9 gene may explain a different phenotype of hereditary oligodontia observed in humans, which affects not only molars but also other tooth lines; and is characterized by tooth small size in both types of dentition. This mutation is characterized by a replacement of the amino acid arginine by tryptophan in a region entirely preserved in all genes of the matched sequenced box [50].
In humans, Pitx2 expression deficiency associated with Rieger syndrome is characterized by oligodontia [51].
9. Conclusion
The biological process is the same for all teeth, including molars, regardless of their identity, but epithelial signaling and homeogenic combination differ from one tooth type to another.
The study of first molar of the mouse has allowed us to better understand and follow the stages of dental development in humans. The general pattern remains the same, unlike the training time, the complexity of the dental system, the presence of two types of teeth in humans, and unlimited incisors growth in mice.
The multidisciplinary approach between fundamental and clinical research is essential to clarify the relationship between molecular involvement and clinical manifestations.
Understanding the molecular mechanisms of dental anomalies, including those affecting human molars, helps to propose diagnostic hypotheses and thus to improve patient management.
Future research should focus on synergizing molecular and genetic approaches to further analyze the action mechanisms of key genes involved in the development of human molars.
Conflict of interest
The authors declare that they have no conflicts of interest with the contents of this article.
\n',keywords:"tooth development, molar, morphological appearance, molecular regulation, epithelial-mesenchymal interaction",chapterPDFUrl:"https://cdn.intechopen.com/pdfs/66644.pdf",chapterXML:"https://mts.intechopen.com/source/xml/66644.xml",downloadPdfUrl:"/chapter/pdf-download/66644",previewPdfUrl:"/chapter/pdf-preview/66644",totalDownloads:126,totalViews:0,totalCrossrefCites:0,dateSubmitted:"December 5th 2018",dateReviewed:"March 7th 2019",datePrePublished:"May 8th 2019",datePublished:null,readingETA:"0",abstract:"Dental development is a complex process by which teeth from embryonic cells grow and erupt into the mouth. It is governed by epithelio-mesenchymal interactions. The biological mechanism is the same for all teeth; however, epithelial signaling and homeogenous combinatorics are different from one type of tooth to another. The primary dental blade splits into the vestibular and primary dental blades opposite to the mesenchymal condensation. During dental development, three successive stages are described: bud, cup, and bell. The secondary dental blade responsible for the formation of germs in permanent teeth is formed from the primary dental blade in the bell stage. For the central incisor, lateral incisor, canine, first temporary molar, and second temporary molar, each primary dental blade gives rise to a single secondary dental blade for the corresponding permanent tooth. On the other hand, the primary dental blade of the second temporary molar will cause the formation of four secondary dental blades that will cause the formation of permanent germs of the second premolar, the first permanent molar, the second permanent molar, and the third permanent molar. The objective of this chapter is to focus on the cellular and molecular mechanisms explaining the normal development of molars by presenting the different current data and theories of science illustrating the human molar embryological development.",reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/66644",risUrl:"/chapter/ris/66644",signatures:"Fatiha Rhrich and Hakima Aghoutan",book:{id:"8837",title:"Human Teeth - Key Skills and Clinical Illustrations",subtitle:null,fullTitle:"Human Teeth - Key Skills and Clinical Illustrations",slug:null,publishedDate:null,bookSignature:"Prof. Zühre Akarslan and Prof. Farid Bourzgui",coverURL:"https://cdn.intechopen.com/books/images_new/8837.jpg",licenceType:"CC BY 3.0",editedByType:null,editors:[{id:"171887",title:"Prof.",name:"Zühre",middleName:null,surname:"Akarslan",slug:"zuhre-akarslan",fullName:"Zühre Akarslan"}],productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"}},authors:null,sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_2",title:"2. Phylogenetic aspects",level:"1"},{id:"sec_3",title:"3. Morphological aspects",level:"1"},{id:"sec_3_2",title:"3.1 Formation of the odontogenic epithelium",level:"2"},{id:"sec_4_2",title:"3.2 Placement of the vestibular and primary dental blades",level:"2"},{id:"sec_5_2",title:"3.3 Evolution of dental placodes",level:"2"},{id:"sec_7",title:"4. Placement of molar dental germs",level:"1"},{id:"sec_8",title:"5. Root formation",level:"1"},{id:"sec_8_2",title:"5.1 Formation of the Hertwig epithelial sheath",level:"2"},{id:"sec_9_2",title:"5.2 Formation of root dentin, cement, and apex",level:"2"},{id:"sec_11",title:"6. Molecular aspects",level:"1"},{id:"sec_11_2",title:"6.1 Epithelial-mesenchymal interactions",level:"2"},{id:"sec_12_2",title:"6.2 Determination of the dental region",level:"2"},{id:"sec_13_2",title:"6.3 Determination of dental identity",level:"2"},{id:"sec_14_2",title:"6.4 Molecular factors involved in root formation",level:"2"},{id:"sec_16",title:"7. Signaling center (primary and secondary enamel knots)",level:"1"},{id:"sec_17",title:"8. Genes and dental problems",level:"1"},{id:"sec_18",title:"9. Conclusion",level:"1"},{id:"sec_22",title:"Conflict of interest",level:"1"}],chapterReferences:[{id:"B1",body:'Pispa J, Thesleff I. Mechanisms of ectodermal organogenesis. Developmental Biology. 2003;262:195-205. DOI: 10.1016/S0012-1606(03)00325-7'},{id:"B2",body:'Tims HWM. The evolution of the teeth in the mammalia. Journal of Anatomy and Physiology. 1903;37(2):131-149. PMC1287046'},{id:"B3",body:'Luo ZX. Transformation and diversification in early mammal evolution. 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Connective Tissue Research. 1995;32:1-4. DOI: 10.3109/03008209509013701'},{id:"B29",body:'Line SRP. Molecular morphogenetic fields in the development of human dentition. Journal of Theoretical Biology. 2001;211:67-75. DOI: 10.1006/jtbi.2001.2333'},{id:"B30",body:'Mitsiadis TA, Smith MM. How do genes make teeth to order through development? Journal of Experimental Zoology. Part B, Molecular and Developmental Evolution. 2006;306(3):177-182. DOI: 10.1002/jez.b.21104'},{id:"B31",body:'Aberg T, Wozney J, Thesleff I. Expression patterns of bone morphogenetic proteins (Bmps) in the developing mouse tooth suggest roles in morphogenesis and cell differentiation. Developmental Dynamics. 1997;210(4):383-396. DOI: 10.1002/(SICI)1097- 0177(199712)210:4<383::AID-AJA3>3.0.CO;2-C'},{id:"B32",body:'Yamashiro T, Tummers M, Thesleff I. Expression of bone morphogenetic proteins and Msx genes during root formation. Journal of Dental Research. 2003;82(3):172-176. 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