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<ul><li><p>REVIEW OF LITERATURE</p><p>15</p><p>REVIEW OF LITERATURE</p><p> The Heteroptera is a numerically large and highly diverse group of insects. </p><p>About 42,300 species belonging to 89 families from this suborder have been described </p><p>many of which are pests of economically important food plants (Henry, 2009).</p><p>Heteroptera comprises eight major infra-orders namely Coleorrhyncha, </p><p>Enlcocephalomorpha, Dipsocoromorpha, Leptopodomorpha, Gerromorpha, Nepomorpha, </p><p>Cimicomorpha and Pentatomomorpha. Within Cimicomorpha, Reduviidae and Miridae </p><p>are the major families. Reduviidae is one of the largest families of terrestrial Heteroptera, </p><p>globally comprising 6601 species and subspecies in 961 genera and 25 subfamilies</p><p>(Maldonado, 1990). These insects are abundant, occur worldwide and are voracious </p><p>predators, thus are named assassin bugs (Ambrose, 1999; Schaefer and Panizzi, 2000). </p><p>Assassin bugs may not be useful as predators of specific pests as they are polyphagous</p><p>but they are valuable predators in situations where a variety of insect pests occur. They </p><p>kill more prey than they need to satiate themselves by their behavior of indiscriminate </p><p>killing. Their importance as predators and their conservation augmention for utilization in </p><p>biocontrol programmes has been discussed by Ambrose (1987, 1999).</p><p>(A) Chromosome complement and meiosis :</p><p>(a) Heteroptera:</p><p>The pioneer contribution towards the cytological aspects of this suborder was made </p><p>by Henking (1891) which was followed by very elaborate cytological investigations by </p><p>Montgomery (1898), McClung (1901) and Wilson (1909 a, b &amp; c). The chromosomes of </p><p>Heteroptera are holocentric and the occurrence of the diffuse centromere was proposed</p><p>and evidenced in a series of publications by Schrader (1935, 1940 a &amp; b, 1947). Schrader </p></li><li><p>REVIEW OF LITERATURE</p><p>16</p><p>(1935) observed chromosomes of Protenor belfragei (Alydidae) to be without a discrete </p><p>localized centromere. Rather spindle fibers were found to be attached all over the surface </p><p>of the chromosome. Further, he observed that the chromosomes showed no bending at </p><p>anaphase-I and two halves remained parallel to one another as they passed on to two</p><p>poles. The hypothesis of diffuse kinetochore was extended to the entire group of </p><p>Heteroptera by Schrader (1940 a &amp; b, 1947). He elucidated the importance of diffuse </p><p>centromere in the evolution of this group. He interpreted that the fragments formed as a </p><p>result of accidental breakage of chromosome with diffuse kinetochore remained </p><p>functional and behaved normally at mitosis and passed through several generations </p><p>thereby changing the fundamental number of a species. Based on chromosome behavior</p><p>during mitosis as well as meiosis, Troedsson (1944), too, claimed that the heteropteran </p><p>chromosomes were holocentric. Hughes-Schrader (1948) examined the behavior of </p><p>heteropteran chromosomes during mitosis and recorded that:</p><p>1. Each chromosome orientated with its long axis at right angle to the polar axis at</p><p>mitotic metaphase.</p><p>2. No primary constriction appeared during mitosis.</p><p>3. Terminalisation of chiasmata was completed by late diakinesis.</p><p>4. Chromosomal fibers were organized along the entire length of each chromatid so </p><p>that mitotic chromatids separate by parallel disjunction.</p><p>These features were further supported by Heizer (1950, 1951), Halkka (1956) and </p><p>Lewis and Scudder (1957). Additionally, holocentricity was demonstrated experimentally </p><p>in the Heteroptera by Hughes-Schrader and Ris (1941) and Ris (1942) also. However, </p><p>Mendes (1949), Parshad (1957 a, b, c &amp; d) and Rao (1958) suggested the heteropteran </p></li><li><p>REVIEW OF LITERATURE</p><p>17</p><p>chromosomes to be monocentric because of the presence of a presumed primary </p><p>constriction at mitotic metaphase. This claim was strongly criticized by Hughes-Schrader </p><p>and Schrader (1961). Piza (1958), on the other hand, suggested that the heteropteran </p><p>chromosome was dicentric. Most of the authors, however, supported the opinion of</p><p>holocentric nature of chromosomes (Manna, 1951; Banerjee, 1959; Wolfe and John, </p><p>1965; Ueshima, 1966, 1979; Ueshima and Ashlock, 1980). La chance et al. (1970) further </p><p>elaborated that even in holocentric chromosomes, some fragments particularly small </p><p>ones, may be eliminated or delayed in the anaphase separation. At the ultrastuctural level,</p><p>Buck (1967) while working on Rhodnius prolixus, observed an extensive layer of </p><p>material all over the mitotic chromosome to which spindle fibers were attached. Meiotic </p><p>chromosomes, on the other hand, did not show this structure. Comings and Okada (1972) </p><p>carried out investigations on formation of kinetochore plates in Heteroptera with a diffuse </p><p>kinetochore during mitosis and meiosis, and discussed their suppression during </p><p>terminalization of chiasmata. They observed a centromere plate extending for upto 75% </p><p>of the length of the mitotic chromosome in Oncopeltus fasciatus but centromeric plate </p><p>was absent in meiotic chromosomes. It was interesting that repeat DNA sequences were </p><p>short and scattered over the chromosome. On the other hand, in organisms with </p><p>monocentric chromosomes, repeated DNA sequences are in high concentration near the </p><p>centromere (Lagowsky et al., 1973). Ruthman and Permantier (1973) found the </p><p>centromere to cover only 4.2% of the entire chromosome in Dysdercus intermedius. </p><p>White (1973) elaborated that the ability of spindle fibres to attach large part of the </p><p>chromosome during cell division becomes the basis for the principle of Karyotypic </p></li><li><p>REVIEW OF LITERATURE</p><p>18</p><p>Orthoselection. The aberrations induced by radiations and chemicals help to understand </p><p>various aspects of the chromosomes, its organization, rearrangements and evolution. </p><p>The behavior of holokinetic chromosomes during the course of mitosis and meiosis </p><p>has been studied by Darlington and Upcott (1938) by taking the measurements of packing </p><p>and contraction in the chromosomes. Darlington (1939) further discussed the genetic and </p><p>mechanical properties of sex chromosomes during division in Heteroptera. The details of </p><p>kinetic activities of autosomes and sex chromosomes during meiosis have been discussed </p><p>by Hughes-Schrader and Schrader (1956, 1957, 1961), Gonzalez-Garcia et al. (1996) and </p><p>Perez et al. (1997, 2000). </p><p>Any scientific study of an organism needs its prior identification, naming and </p><p>classification. Taxonomists use various morphological characters to bring different </p><p>members of a group of plants and animals under a systematic list showing their natural </p><p>phylogenetic relationships. With the growing knowledge of nuclear structures, the </p><p>cytologists believe that chromosomal attributes are also morphological and can be </p><p>substantially utilized for the purpose of taxonomy in addition to general morphological </p><p>features. In Heteroptera, karyotype is prone to evolutionary changes either by fusions or </p><p>fragmentations of chromosomes which lead to decrease or increase in the chromosome </p><p>number in different species. Every fusion and fragmentation appears first in a </p><p>polymorphic state prior to its fixation in the karyotype. The importance of heteropteran </p><p>material for cytological studies from evolutionary point of view was recognized in the </p><p>beginning of twentieth century by Montgomery (1901a &amp; b, 1904, 1906), Wilson (1905a, </p><p>b &amp; c, 1906, 1907a, b &amp; c, 1909a &amp; b, 1910, 1911, 1912, 1913, 1932) and Browne (1916) </p><p>who carried out cytological investigations along with morphological characters of </p></li><li><p>REVIEW OF LITERATURE</p><p>19</p><p>Heteroptera. Payne (1909, 1910, 1912), Chickering (1927a &amp; b, 1932, 1933) and </p><p>Nishimura (1927) stressed upon the importance of chromosome studies of different </p><p>families of Heteroptera with respect to their karyotypic evolution. This practice is </p><p>becoming more and more obvious in modern systematics. Cytologists are interested in </p><p>collecting the cytological data of more and more species of Heteroptera. At international </p><p>level, karyological studies on Heteroptera have been done by Foot and Strobell (1907, </p><p>1912, 1914), Schrader (1932, 1935, 1939, 1940a &amp; b, 1941a &amp; b, 1945a &amp; b, 1946a &amp; b, </p><p>1947, 1960a &amp; b), Toshioka (1933, 1934, 1935, 1936, 1937), Geitler (1937, 1938, 1939a </p><p>&amp; b), Pfaler-Collander (1937, 1941), Freeman (1940), Mc Clung (1940), Hughes-</p><p>Schrader (1931, 1940, 1942, 1948, 1955), Schrader and Hughes-Schrader (1956, 1958), </p><p>Ekblom (1941), Troedsson (1944), Yosida (1944, 1946, 1947, 1950, 1956), Xavier </p><p>(1945), White (1948), Heizer (1950), Lewis and Scudder (1957), Leston (1957, 1958), </p><p>Ueshima (1963, 1966, 1979), Mikolajski (1964, 1965, 1967a &amp; b, 1968, 1970,1971),</p><p>Takenouchi and Muramoto (1964, 1967, 1968, 1969, 1970a &amp; b, 1971a &amp; b, 1972a &amp; b, </p><p>1973), Muramoto (1973a, b, c &amp; d, 1974, 1975a, b &amp; c, 1976, 1977, 1978a &amp; b, 1979, </p><p>1981, 1982, 1985), Akingohungbe (1974), Kuznetsova and Petropavlovskaya (1976) and </p><p>Reddi and Chari (1976, 1978).</p><p>Extensive cytological studies have been carried out on Heteroptera by Ueshima </p><p>and Ashlock (1980), Sands (1982a &amp; b), Nuamah (1982), Newman and Cheng (1983), </p><p>Nokkala and Nokkala (1983, 1984, 1997, 1999, 2004), Papeschi and Bidau (1985), </p><p>Nokkala (1985, 1986). Papeschi (1988, 1991, 1992, 1994, 1995, 1996), Papeschi and </p><p>Mola (1990 a &amp; b), Grozeva and Kuznetsova (1990, 1993), Panzera et al. (1992, 1995, </p><p>1996, 1997, 1998, 2000), Perez et al. (1992, 1997, 2000, 2004, 2005), Grozeva (1995a &amp; </p></li><li><p>REVIEW OF LITERATURE</p><p>20</p><p>b, 1997), Nokkala and Grozeva (1997), Bressa et al. (1998, 1999, 2001a &amp; b, 2002, 2003, </p><p>2005, 2008), Rebagliati et al. (1998, 2001, 2002, 2003, 2005, 2010 a &amp; b), Tartarotti and </p><p>Azeredo-Oliveira (1999a &amp; b), Jacobs and Liebenberg (2001), Grozeva and Nokkala </p><p>(2002), Jacobs and Groenveld (2002), Papeschi and Bressa (2002, 2004, 2006), Papeschi</p><p>et al. (2003), Severi-Aguiar and Azeredo-Oliveira (2003, 2005), Nokkala et al. (2003, </p><p>2006 a &amp; b), Jacobs (2004), Grasiela et al. (2004), Ituarte and Papeschi (2004), Kerzhner </p><p>et al. (2004), Franko (2005), Grozeva et al. (2005, 2006, 2007, 2008, 2009), Franko et al. </p><p>(2006), Bressa and Papeschi (2007), Poggio et al. (2007), Souza et al. (2007a, b &amp; c, </p><p>2008 a &amp; b, 2009, 2010), Bardella et al. (2008), Kuznetsova and Grozeva (2008), Pires </p><p>(2008), Toscani et al. (2008), Zhang and Zheng (1999), Kaur and Semahagn (2010 a) </p><p>and Yang et al. (2012).</p><p>In India, major contribution to the cytology of Heteroptera has been made by </p><p>Manna (1950, 1951, 1957, 1958, 1962, 1982, 1983, 1984) describing the chromosome </p><p>complements of several Indian species of different families of Heteroptera and discussing</p><p>cytoevolutionary trends. The course of meiosis and chromosomal behaviour in various </p><p>heteropterans species belonging to different families have been discussed by Das Gupta </p><p>(1950), Ray Chaudhuri and Manna (1952, 1956), Bawa (1953), Rao (1954, 1955, 1958), </p><p>Dutt (1955, 1957), Sharma and Parshad (1955a &amp; b, 1956), Das (1956, 1958), Parshad </p><p>(1956, 1957a, b, c &amp; d, 1958), Sharma et al. (1957), Srivastava (1957, 1965), Banerjee </p><p>(1958, 1959), Bagga (1959), Ray Chaudhuri and Banerjee (1959) and Jande (1959 a, b &amp; </p><p>c, 1960 a, b &amp; c). Manna (1962, 1982, 1983, 1984), Rajasekharasetty (1963), </p><p>Bhattacharya and Halder (1978), Malipatil (1979), Manna and Deb-Mallick (1980, 1981a </p><p>&amp; b, 1982, 1983, 1984), Barik et al. (1981), Manna and Dey (1981), Mittal and Joseph </p></li><li><p>REVIEW OF LITERATURE</p><p>21</p><p>(1981, 1982, 1984), Manna et al. (1985), Dey and Wangdi (1985, 1988, 1990), Satapathy </p><p>and Patnaik (1988, 1989, 1991), Satapathy et al. (1990), Kaur et al. (2006, 2009), Kaur </p><p>and Suman (2009), Kaur and Singh (2010) and Kaur and Semahagn (2010 b) made a </p><p>significant contribution towards meiotic studies in Heteroptera.</p><p>Heteroptera is characterized by sex determination mechanisms of simple </p><p>(XY/XX, XO/XX), multiple (XnO/XnXn, XnY/XnXn and XnYn/XX) and neo-XY types </p><p>(Chickering and Bacorn, 1933; Schrader, 1940a &amp; b; Manna, 1951, 1984; Jande, 1959b </p><p>&amp; c; Ueshima, 1979; Nokkala and Nokkala, 1983; Nokkala et al., 2003). Different views </p><p>have been proposed about the origin and evolution of sex determining systems in </p><p>Heteroptera. Ueshima (1979) considered XO system encountered commonly in primitive </p><p>heteropteran taxa to be the ancestral one and XY to be derived from it. It was further </p><p>suggested that multiple sex chromosome systems of Heteroptera evolved from the XY </p><p>system. By contrast, Nokkala and Nokkala (1983, 1984) considered XY system to be the </p><p>ancestral one and dominant sex mechanism and XO system to be derived from it. In </p><p>Heteroptera, multiple sex mechanisms are not accompanied by any reduction in autosome </p><p>number implying their origin by fragmentation of sex chromosomes. Evolution of </p><p>multiple sex determination mechanisms have been discussed by Payne (1909), Troedsson </p><p>(1944), Ueshima (1979), Barik et al. (1981), Manna (1982, 1984), Grozeva and Nokkala </p><p>(1996) and Bressa et al. (1999). Neo-XY sex determination system is thought to be </p><p>originated as a result of autosome-sex chromosome fusion. Involvement of autosomes in </p><p>sex mechanisms has been discussed by Chickering and Bacorn (1933), Schrader (1940a </p><p>&amp; b), Jande (1959b &amp; c), Ueshima (1979), Bressa et al. (1999, 2009), Nokkala and </p></li><li><p>REVIEW OF LITERATURE</p><p>22</p><p>Nokkala (1999), Jacobs (2003, 2004), Papeschi and Bressa (2006) and Poggio et al. </p><p>(2007). </p><p>In groups with holocentric chromosomes, meiosis is post-reductional i.e. </p><p>chromosomes divide equationally during first meiotic division and reductionally during </p><p>the second meiotic division. This is true of Odonata, Trichoptera and Lepidoptera.</p><p>Heteropterans are unique in showing pre-reductional meiosis for autosomes and post-</p><p>reductional for the sex chromosomes. In Heteroptera, chiasma frequency is low. Single </p><p>chiasma per bivalent is the predominant condition. Sex chromosomes remain </p><p>achiasmatic. The general course of meiosis, behavior of different types of chromosomes </p><p>(autosomes, m-chromosomes, sex chromosomes) during meiosis and chiasma frequency </p><p>have been described and discussed by Das Gupta (1950), Ray Chaudhuri and Manna </p><p>(1952, 1955), Battaglia and Boyes (1955), Dutt (1955, 1957), Manna (1962, 1982, 1983, </p><p>1984), Takenouchi and Muramoto (1964, 1967, 1968, 1970a &amp; b, 1971a &amp; b, 1972a &amp; </p><p>b), Ueshima (1966, 1979), Muramoto (1973a, b, c &amp; d, 1974, 1975a, b &amp; c, 1978 b, </p><p>1979), Camacho et al. (1985), Mola and Papeschi (1993), Bressa et al. (1998, 2001a &amp; </p><p>b), Jacobs and Liebenberg (2001), Rebagliati et al. (2001, 2003), Jacobs and Groeneveld </p><p>(2002), Bressa et al. (2002), Papeschi et al. (2003), Poggio et al. (2007, 2011) and Kaur </p><p>et al. (2010). </p><p>Heteroptera is characterized by the presence of a pair of minute elements in some </p><p>families. The term m-chromosomes was first assigned by Wilson (1905 b) to a pair of </p><p>very small chromosomes in Coreid bugs which differed in meiotic behavior from both </p><p>autosomes and sex chromosomes. They remained unpaired or achiasmatic during meiotic </p><p>prophase and associated terminally to form a pseudobivalent at metaphase-I to ensure </p></li><li><p>REVIEW OF LITERATURE</p><p>23</p><p>regular segregation. Wilson (1905b) termed this pairing near metaphase-I to as touch </p><p>a...</p></li></ul>

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