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Author(s): Jian-Ying Guo , Gang Wu , and Fang-Hao Wan Source: Journal of Economic Entomology, 106(3):1379-1385. 2013. Published By: Entomological Society of America URL: http://www.bioone.org/doi/full/10.1603/EC12401
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In China. also owing to its expanding host range (Tsueda and Tsuchida 2011) and its ability to reach high population densities (Henneberry et al. the B. (2006) demonstrated that host plants can mediate the success of population establishment of B.1. College of Plant Sciences and Technology. tabaci MEAM1 cryptic species (Zang et al. Rate of increase was also higher in the second generation. Haidian. Institute of Plant Protection. the survival. and reproduction of Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae) MEAM1 cryptic species were compared over two consecutive generations on three cotton cultivars of different gossypol levels. Both cotton cultivar and generation signiÞcantly affected the Þtness of the whiteßy. The comparison of the life histories of B. the putative Middle EastÐAsia Minor 1 species (MEAM1) (previously known as biotype B. 2009). Meng et al. most provinces of China henceforth (Zhang 2000. It causes worldwide attention. De Barro et al. In this study. (1999) found a positive correlation between gossypol level and the population abundance of A. 1993. Beijing 100081. (1964) reported that the infestation of cotton aphid. In both generations. primarily begomoviruses (Perring et al. Entomol. It can cause damage directly by feeding and indirectly by transmission of plant pathogenic viruses.org/10. 1995) and to tolerate high temperatures (Mun ˜ iz and Nombela 2001). On each cultivar. gossypii. Plant secondary substances are an important biochemical basis for plant resistance to insects and are desirable tools in most pest management programs by complementing the actions of insecticides and natural enemies (Ascher 1993. Wuhan 430070.3 AND FANG-HAO WAN1 J. 2011). Aphis gossypii Glover (Homoptera: Aphididae). Tsueda and Tsuchida 2011). 2005). Gao et al. China. tabaci MEAM1 cryptic species in Australia. Cuthbertson et al. Huazhong Agricultural University. Econ.issg. the immature development time was shorter and the immature survival rate was higher in the second generation than those in the Þrst generation. Host plant is an important factor affecting the biology and population dynamics of B. Liu et al. Zhong-Guan-Cun Nan-Da-Jie. DOI: http://dx. China. #12. Shi-Zi-Shan St. a phenolic sesquiterpenoid aldehyde. Hongshan.. increased in cotton cultivars lacking the gossypolproducing glands. as reviewed by Drost et al. the immature development times on the low-gossypol cultivar ZMS13 were signiÞcantly longer than those on the high-gossypol cultivar M9101 or medium-gossypol cultivar HZ401.HORTICULTURAL ENTOMOLOGY Effects of High-Gossypol Cotton on the Development and Reproduction of Bemisia tabaci (Hemiptera: Aleyrodidae) MEAM1 Cryptic Species JIAN-YING GUO. KEY WORDS Bemisia tabaci gossypol. development The whiteßy Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae) is a cryptic species complex that contains Ͼ28 morphologically indistinguishable species (De Barro et al. Greenberg et al. tabaci MEAM1 cryptic species on the low-gossypol cotton cultivar ZMS13 was higher than that on the medium. 2012). tabaci MEAM1 cryptic species on different cotton varieties is important for the development of an integrated pest management program of the whiteßy by using plant secondary metabolic compounds. These results demonstrated that the Þtness of B. 106(3): 1379Ð1385 (2013).org/database). hereon MEAM1) has become one of the most globally destructive pests of vegetable crops (Global Invasive Species Database. Gossypol. The female fecundity and rate of population increase of the whiteßy ranked in the following order: ZMS13 Ͼ HZ401 Ͼ M9101. Bottger et al. survival rate. e-mail: guojianying@caas. 2011. 2010). 2 Corresponding author. tabaci MEAM1 cryptic species. 2006.2 GANG WU. 3 Department of Plant Protection.cn. 2011. (1998). These studies support the general assumption that the broad host plant adaptation ability plays an important role in the successful invasion process of B. 2007.00/0 ᭧ 2013 Entomological Society of America . #1. (2008) indicated that high levels of gossypol in cotton lowered the adult 0022-0493/13/1379Ð1385$04.1603/EC12401 ABSTRACT Use of plant secondary metabolic compounds is an important method for insect pest control. Within this whiteßy complex. Liu et al. Delatte et al. is an important allelochemical produced by the subepidermal glands of cotton cultivars and exhibits antibiosis against some cotton pests (Wu et al. development.doi. tabaci MEAM1 whiteßy Þrst was found in the late 1990s and outbreaks in 1 State Key Laboratory for Biology of Plant Diseases and Insect Pests.or high-gossypol cultivar. Chinese Academy of Agricultural Sciences. http://www. Hu et al.
and those that did not hatch by day 9 were considered dead and removed. and available potassium of 262. Adult sex was determined with the aid of a stereomicroscope. and one egg on each plant was left for observation. the leaf clip-on cage was removed and the plant was checked under the stereomicroscope. it is difÞcult to draw conclusions on the effect of gossypol on B. RH. But until now. tabaci adults (Ͻ12 h old) were collected from the colony and paired with the aid of a stereomicroscope. After 8 h.7 mg kgϪ1 (determined by hydraulic N. tabaci observed on nongossypol cultivars. 106. As a major cotton pest. During the 8-to-10-leaf stage of cotton (Ϸ40 Ð50 d after planting). and M9101. gossypii. ZMS13. and each pair of whiteßies was conÞned on the lower leaf surface of a fresh cotton plant as its immature stage by using a leaf clip-on cage. the plants were checked under a stereomicroscope (126ϫ magniÞcation) to conÞrm that they were not infested by insects. as described above. (2010) reported signiÞcantly lower relative growth rates of beet armyworm.0 mg kgϪ1 (titration with 0. (2008) revealed high gossypol concentrations in artiÞcial diet reduced the survival rate and pupal weight of Helicoverpa virescens larvae. There was only one clip-on cage per plant. These pots with cotton or tomato seedlings were placed randomly in the insectary and rerandomized once a week to minimize position effects. Each pair of whiteßies (one adult female and one adult male) was conÞned on the lower leaf surface of a cotton plant by using a leaf clip-on cage (Zang et al. and photoperiod conditions as for the whiteßy colony maintenance. ZMS13. Stipanovic et al. 2005). Ozgur and Sekeroglu (1986) reported high numbers of B. 8 cm in height) in an insectary at the Experimental Station. tabaci were recorded daily until adult emergence. The plants were checked daily to determine the development of the whiteßies. thrips.20% racemic gossypol were 4. These cotton and tomato seeds were grown individually in plastic pots (9 cm in diameter. Each treatment was repeated three times at an interval of 2 wk.70 Ϯ 10% RH. (Solanaceae) ÔGoldensunÕ were obtained from the Yigao Seed Company. tabaci MEAM1 cryptic species. and the whiteßy eggs were counted with the aid of a stereomicroscope. Because high-gossypol cotton cultivars are resistant to various insect pests. 60 eggs were used for each cotton cultivar treatment. New plants were used as necessary. they have been widely adopted for pest management by cotton growers. Unhatched eggs were marked on day 6. Langfang. Newly emerged adults were collected from each cotton cultivar treatment and counted daily. no. . were obtained from the Institute of Cotton Research. Henan province. 15 pairs were used for each cotton cultivar treatment. tabaci MEAM1 Cryptic Species.5 M NaHCO3 solution). Wu et al. 3 survival and fecundity and prolonged nymphal duration of A. China. Institute of Plant Protection. and aphids. Hebei Province.44.1. Spodoptera exigua (Hu ¨ bner) (Lepidoptera: Noctuidae). development. the remaining eggs were removed carefully by using an artistÕs Þne brush (size 000). HZ401. In total. with organic matter availability of 14. survival. tabaci MEAM1 cryptic species colony was reared on potted tomato plants in another insectary at the Langfang Experimental Station.. The colony was maintained for over 10 generations before the start of the experiment. The plant bearing one whiteßy egg was transferred to a clean cage and moved into an insectary of the same temperature. CAAS. Schuster (1979) found that high-gossypol cotton was susceptible to B. tabaci on different cotton varieties and thereafter develop ecologically sound integrated pest management (IPM) programs (Mun ˜ iz and Nombela 2001). available nitrogen of 397. CAAS. Adults of each treatment were paired. The B. Lukefahr and Houghtaling (1969) found that high-gossypol cotton cultivar signiÞcantly reduced the population abundance of tobacco budworms. Stipanovic et al. 0. 70 Ϯ 10% RH. ZMS13. Five pairs of whiteßies were used with three replications for each cotton cultivar treatment (i. with gossypol content of 0.2 mg kgϪ1 (titration with 1N CH3COONH4 solution).. Materials and Methods Host Plants. or four-to-Þve-leaf stage of tomato (Ϸ25Ð30 d after planting). Heliothis virescens (F. The whiteßies were transferred to a new leaf daily. The development and survival of individual B. Hebei province. Chinese Academy of Agricultural Sciences (CAAS). and reproduction of the whiteßy on three cotton cultivars with different gossypol levels were compared over two consecutive generations. under the conditions of 27 Ϯ 1ЊC. In total.) (Lepidoptera: Noctuidae). and a photoperiod of 14:10 (L:D) h.5%. and a photoperiod of 14:10 (L:D) h. at 27 Ϯ 2ЊC. Institute of Plant Protection. The soil had a pH of 7. tabaci. and M9101). HZ401. Anyang. Seeds of tomato. On the contrary. respectively (Gao et al.8 times higher than those on a diet containing 0. we hypothesized that high-gossypol cotton decreases the Þtness of B. Newly emerged B. (Malvaceae) cvs. HZ401.24% racemic gossypol.1380 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. In this study. No further fertilizer or insecticide was used during the experiment. Lycopersicon lycopersicum L. and M9101). mites.e. available phosphorus of 269. China. tabaci. Life History Parameters of B. A group of four plants with whiteßies was placed in a cubic cage (60 cm on each side) covered with a Þne mesh net to prevent the incursion of other insects. tabaci MEAM1. Twenty pots were used in each of the three cotton cultivar treatments (i. The pots were watered every 3 d after cotton seedling emergence. 1 N NaOH hydrolysis).12%. Insects. Gossypium hirsutum L. it is essential to understand the biology of B. These uninfested plants were used for the experiment.06. China. 2008).e. such as whiteßies. and 1. To determine the effects of gossypol on the Þtness of the B. (2006) reported that the survival rates of Helicoverpa zea (Boddie) (Lepidoptera: Noctuidae) larvae on a diet containing 0. The plants were kept in the same insectary used for the observation of the development of the immature whiteßies. Seeds of cotton. Langfang. feeding on high-gossypol cotton cultivar in three consecutive generations.
fourth-instar duration.0a HZ401 83.05) and total amino acid (P Ͼ 0. Results Foliar Chemical Composition Assays of Cotton Plants. However. SigniÞcantly lower foliar nitrogen (P Ͻ 0. survival. Multi-way ANOVAs of Cotton Cultivar and Generation Effects on B. There were also no signiÞcant higher-order interactions between replication or sex and cotton cultivar or generation. Europa Elemental Instruments. tabaci were not affected by replication (four-way ANOVA: egg duration.001) were also signiÞcantly affected by cotton cultivar. tabaci TO GOSSYPOL 1381 The process was continued until the death of all of the individuals. Nitrogen content was measured using a CNH analyzer (model ANCA-nt.001). During the adult emergence peak.2 Ϯ 2.9a 90. Okehampton.004 0. immature development time. as a proportion of fresh weight.05b 4.09 Ϯ 0.05) higher-order interaction terms. and those on the number of eggs square roottransformed before analysis (Sokal and Rohlf 1995). and then preserved in a refrigerator at Ϫ20ЊC for subsequent chemical composition assays. nymphal duration.1a ZMS13 84.06a 4. P ϭ 0. second-instar duration. where NtϪ1 is the number of the initial population and Nt is the number of the future population (Southwood and Henderson 2000). The interaction of cotton cultivar and generation also signiÞcantly affected the immature survival rate (P Ͻ 0.7a P 0.127. P ϭ 0. immature duration. generation.324.462. Co. immature survival rate. These data between generations were compared by the StudentÕs t-test. was calculated after drying the leaves at 80ЊC for 72 h. the population is decreasing. tabaci MEAM1 Cryptic Species. Comparable leaves with those in the whiteßy life history studies were collected randomly from extra cotton plants at the beginning of the Þrst-generation rearing. Data on the adult longevity and number of eggs were compared by three-way ANOVA with cotton cultivar. P ϭ 0. Þrst-instar duration. and replication as factors.789. foliar water (P Ͼ 0.11 Ϯ 0. P ϭ 0.539. Therefore. China.05) contents did not differ signiÞcantly among these cotton cultivars (Table 1). P ϭ 0.001) and fecundity (P Ͻ 0. nitrogen content.51 Ϯ 0.3 Ϯ 1. third-instar duration.001 Mean Ϯ SD.01b 0. P ϭ 0. Data Analysis. protein (P Ͻ 0. P ϭ 0.01b 0.05). P ϭ 0. If R Ͼ 1.01a 88. and free fatty acid (P Ͻ 0. egg hatching rate.: RESPONSE OF B. Data on the immature survival rate and percentage of female adults were compared by two-way ANOVA among treatments (type III). The longevity of individual B. and male adult longevity of B.7 Ϯ 2. placed in liquid nitrogen for 3 h.com).njjcbio.001) contents were detected in the high-gossypol cotton cultivar M9101 than those in the other two low-gossypol cultivars ZMS13 and HZ401. All observed data were tested for homogeneity of variance by BartlettÕs test and for outliers by DixonÕs test (Sokal and Rohlf 1995). These data among cotton cultivars were compared by the least signiÞcant difference test after one-way ANOVA. Different lowercase letters within a row indicate signiÞcant differences among the three cotton varieties (ANOVA: least signiÞcant difference (LSD) test at P Ͻ 0. In this study.870. total amino acid. female percentage (P Ͻ 0. P ϭ 0.944.05).24 Ϯ 0.105 Ͻ0. Þrst-instar duration. and percentage of female adults were arcsine-transformed. Nanjing.3 Ϯ 2. Protein. data were pooled and compared among cotton cultivars and generations only. Leaf water content. generation.243 4.001). During the treatment of Þrst generation.June 2013 GUO ET AL.001). P ϭ 0. immature duration. P ϭ 0.5 statistical package (Lu 2002).505). Data on the adult fecundity were also not affected by replication for any parameter (three-way ANOVA: male adult longevity. www. All calculations were done using the SPSS 11.9 Ϯ 1. P ϭ 0. P ϭ 0. and free fatty acid contents were assayed according to the directions of the Reagent Kits (Nanjing Jiancheng Ltd. Multifactorial statistical models were reduced by stepwise deletion of nonsigniÞcant (P Ͼ 0. United Kingdom). sex.307) or by sex (four-way ANOVA: egg duration. Female adult longevity (P ϭ 0. Data on the water content. R is calculated as R ϭ immature survival rate (%) ϫ Table 1.3 Ϯ 2. the population is assumed to be increasing and expanding.10 Ϯ 0.268. Both cotton cultivar and generation signiÞcantly affected the immature survival rate. with cotton cultivar and generation as factors.909. those on the immature development time and adult longevity log-transformed.07a Ͻ0.671).558. The index of rate of increase is calculated as R ϭ Nt/NtϪ1. and replication as factors. adults were collected from these extra plants and used for the life history study of the second generation. Jiangsu province. Water and nutrient contents of the three experimental cotton cultivars Indices Water content (%) Nitrogen content (%) Protein (g/liter) Total amino acid (mol/ml) Free fatty acid (mol/ml) Cotton cultivars M9101 82. Five leaves with three replicates were taken from each cotton cultivar. and reproduction of the whiteßy were tested using the same procedure as described for the Þrst generation.902. female adult longevity. Data on the immature development time were compared by four-way analyses of variance (ANOVAs) with cotton cultivar. P ϭ 0.0a 90.3a 219 Ϯ 3c 230 Ϯ 5b 235 Ϯ 4a 0.43 Ϯ 0. P ϭ 0. second-instar duration.292. third-instar duration. percentage of female adults (%) ϫ number of eggs laid per female. nymphal duration. tabaci adult and the female fecundity were determined. tabaci (two-way ANOVA: P Ͻ 0. Development. 120 extra pots with B. fourth-instar duration..001 0.552. number of eggs per female.001). P ϭ 0. Data on the immature development of B. P ϭ 0. tabaci eggs (one egg per pot prepared as described in the previous section) were set up for each cotton cultivar treatment. female adult longevity . and if R Ͻ 1.073. Foliar Chemical Compositions Assays of Cotton Plants.
5 Ϯ 0.8bA 0.001 Ͻ0.001*** 0.001).0bB 3.021) (Table 3).024 6.6 Ϯ 0.001 Ͻ0.0bB 21.113 0. and on M9101 did not differ signiÞcantly (Table 4).021 Ͻ0.001 Cultivara 0.010* 0.693 0.028 0.0 Ϯ 1. In the Þrst generation. on ZMS13 was signiÞcantly higher in the second generation.052 0.5 Ϯ 2. which mainly happened in the Þrst-instar nymphal stage (P Ͻ 0.0bB 7.7aA Ͻ0.7aA 2. and the difference was signiÞcant on ZMS13 (P ϭ 0.4 Ϯ 1.6 Ϯ 1. data were pooled to compare among the three cotton cultivars at a deÞned generation by one-way ANOVA.5aB 0. In the Þrst generation.2 Ϯ 1.087 Ͻ0. On each cotton cultivar. mainly happened in the second-instar nymphal stage (Table 4).001 Ͻ0.001*** 0.001 P Ͻ0.9bB 19.0aB Ͻ0. Different lowercase letters indicate signiÞcant differences among the three cotton cultivars at a deÞned generation (ANOVA: LSD test at P Ͻ 0.8bA 22. Whiteßy generation (the Þrst and second generation of B.4 Ϯ 1.5 Ϯ 1.001** Ͻ0. On each cotton cultivar. In the second generation. 106.7 Ϯ 0.0 Ϯ 0.001 0.131 6.048* 0. All tested eggs hatched on the three cotton cultivars and in both generations.632 0. **.008 2. The nymphal duration was also shortened in the second generation to some extent.001 Second generation 5. In the Þrst generation. P Ͻ 0. the immature survival rates of the whiteßy did not differ signiÞcantly among these cotton cultivars (P ϭ 0. the male longevities did not differ signiÞcantly between the two generations (Table 5).048).4 Ϯ 0.001** 0.05.5 Ϯ 0.0 Ϯ 1. *.243 0. the female percentage on HZ401 was signiÞcantly lower in the second generation.7aA 7.8bA 3.8aA 3.4bA 16.4 Ϯ 0.001 14.4 Ϯ 0.474 0.01.3bA 23.153).001.103 Ͻ0.001).8 Ϯ 0.7bA 7.001*** Ͻ0. respectively. In the second generation.3bA 14.020* First-instar duration (d) Second-instar duration (d) Third-instar duration (d) Fourth-instar duration (d) Nymphal duration (d) Immature duration (d) Cotton cultivars (M9101.001*** 0. The nymphal and immature durations of the whiteßy on the low-gossypol cultivar ZMS13 were signiÞcantly longer than those on the high-gossypol cultivar M9101 or medium-gossypol cultivar HZ401 (P Ͻ 0. In the second generation.4 Ϯ 0. Immature Survival Rates.4bA 15.5 Ϯ 0.990 0.5aB Ͻ0.4 Ϯ 0.9bA 6.001 22.0aA 7.05).212 Ͻ0.001*** Ͻ0.001*** 0.189 0.621 and 0.019).010* 0.4 Ϯ 1. which was mainly caused by the prolonged development times in the second. . Immature Development Times.789 0.608 0.05).562 0. the male adult longevities of the whiteßy did not differ signiÞcantly among the three cotton cultivars (P ϭ 0. and P Ͻ 0.8aA 3. and fecundity (P ϭ 0.4 Ϯ 0.041* 0.001).6aA Ͻ0.001*** Ͻ0.270 Ͻ0.0 Ϯ 0.004** 0.210 0.2aB 5.051 2. the immature survival rate on the low-gossypol cultivar ZMS13 was signiÞcantly higher than that on the high-gossypol cultivar M9101 or medium-gossypol cultivar HZ401 (P Ͻ 0.798 0.2 Ϯ 1.099 0. The differences Mean Ϯ SD. the male longevity on ZMS13 was signiÞcantly longer than that on HZ401 or M9101 (P ϭ 0. the egg and immature durations of the whiteßy were signiÞcantly shorter in the second generation than those in the Þrst generation (P Ͻ 0.9aA Ͻ0.7aA 3.5bA 2.020) (Table 2).001*** 0. tabaci MEAM1 cryptic species Indices Egg duration (d) Host plant M9101 HZ401 ZMS13 P M9101 HZ401 ZMS13 P M9101 HZ401 ZMS13 P M9101 HZ401 ZMS13 P M9101 HZ401 ZMS13 P M9101 HZ401 ZMS13 P M9101 HZ401 ZMS13 P First generation 7.2 Ϯ 1. female percentages did not differ signiÞcantly among these cotton cultivars (P ϭ 0.001) (Table 3).112).094 0. tabaci in both generations (P Ͻ 0. HZ401.001) (Table 4). 3 Table 2. or to compare between the two generations on a deÞned cotton cultivar by StudentÕs t-test. tabaci MEAM1 cryptic species). Two-way ANOVA of the development and reproduction of Bemisia tabaci MEAM1 cryptic species on different cotton cultivars in two consecutive generations Source of variation Egg duration First-instar duration Second-instar duration Third-instar duration Fourth-instar duration Nymphal duration Immature duration First-instar survival Second-instar survival Third-instar survival Fourth-instar survival Immature survival Female percentage Male adult longevity Female adult longevity Female fecundity a b Table 3.001 2.001*** Generationb Ͻ0.004 0.4bA 15.0aB 5. Comparing the two generations on the same cotton cultivar.001 0. Different uppercase letters indicate signiÞcant differences between the two generations on a deÞned cotton cultivar (StudentÕs t-test at P Ͻ 0. Cotton cultivar signiÞcantly affected the nymphal duration of B.6aA Ͻ0.7 Ϯ 1. and *** indicate signiÞcant difference at P Ͻ 0.002** 0. Longevities of Adults.027 0.173 0.441 0. the immature survival rate was signiÞcantly higher in the second generation than that in the Þrst generation. The differences of female longevities among the three cotton cultivars and between the two generations were similar to these of the male longevities (Table 5).8aA 0.057 0.1382 JOURNAL OF ECONOMIC ENTOMOLOGY Vol.007** Ͻ0.1 Ϯ 2.001** 0.001** Ͻ0.8 Ϯ 1.392 0.6 Ϯ 0. (P ϭ 0.123 0.6bA 2.6aA 0.and fourth-instar nymphal stages (P Ͻ 0.3 Ϯ 1.955 Cultivar ϫ generation 0.1aB Ͻ0.9 Ϯ 0. Because of such interactions.8 Ϯ 0.9 Ϯ 0.001 Ͻ0.231 0.001*** 0.6bA 0.6aB 2.775 Ͻ0. respectively).0 Ϯ 1. and ZMS13).1 Ϯ 0.001 19.2 Ϯ 0.452 3. Percentages of Female Adults.001*** 0.6bA 1.7aA 2.001).370 0. the female percentage of B.2 Ϯ 1.017* 0.001 14. tabaci on HZ401 was signiÞcantly higher than that on the other two experimental cotton cultivars (P ϭ 0.050) (Table 4).763 0.2 Ϯ 1.3 Ϯ 0.101 0.791 0.621 3. although it did not affect the egg duration in either generation (P ϭ 0.7bA 2.0 Ϯ 0. Effects of cotton cultivar on the development of immature B.437 0.001 2.6cA 7.8aA Ͻ0.7aA 0.001*** 0.598 Ͻ0. no.2 Ϯ 0. On each cotton cultivar.0 Ϯ 1.452.008).
1aB 98.4bA 21.28 72. virescens (Lukefahr and Houghtaling 1969.132 M9101 100. R ϭ immature survival rate (%) ͓times͔ percentage of female adults (%) ͓times͔ number of eggs laid per female.174 P 0.7aA 97. Although the whiteßy cryptic species in the study of Schuster (1979) is not easy to determine.1aA 53.019 ZMS13 48.8 Ϯ 2.0 Ϯ 0.0bA 19. Rates of Population Increase.0 Ϯ 5.932 0. we speculate that the whiteßy studied by Ozgur and Sekeroglu (1986) is Asia 1 cryptic species.2 Ϯ 9.3 Ϯ 2.082 0. These results conÞrmed our hypothesis that the Þtness of B.001 25.1 Ϯ 1.0aA 100. Here. and environment-friendly method for pest control (Sharma and Ortiz 2002). 2010).Gao et al.046 P Ͻ0.037 (%) ZMS13 76.0 Ϯ 0. In this study.14 31.3 Ϯ 4. the better performance of the whiteßy on ZMS13 might be an integrated effect of low gossypol and high nutrient contents.1aA 0.223 survival rate HZ401 79. S.3 Ϯ 15. 2008).001 0.268 Fourth-instar M9101 96.4 Ϯ 1. tabaci MEAM1 cryptic species decreases when rearing on high-gossypol cotton.3aA 0.699 0. These differences were more dramatic in the second generation. The adverse impacts were more dramatic on the highgossypol cultivar M9101.18 41.730 0.374 P 0.421 Ͻ0.7 Ϯ 3. host plant effect has been identiÞed as an important factor affecting its Þtness (Drost et al. Host plant resistance to insects is an effective.4aA 0.1aB 154.5 Ϯ 3.3aA 83. H. tabaci MEAM1 cryptic species Indices First-instar survival rate (%) Host plant First generation Second generation P M9101 56.071 0.3 Ϯ 10.4aA 0. In both generations.7 Ϯ 7. Effects of cotton cultivar on the immature survival and adult sex ratio of B. 2009).3aA 0. of eggs laid per female Rate of pop increase (R)a Host plant M9101 HZ401 ZMS13 P M9101 HZ401 ZMS13 P M9101 HZ401 ZMS13 P M9101 HZ401 ZMS13 First generation Second generation P 0. higher on ZMS13. tabaci MEAM1 cryptic species are accordant with that of Ozgur and Sekeroglu (1986).4 Ϯ 15. the rate of increase ranked in the following order: ZMS13 Ͼ HZ401 Ͼ M9101.422 Immature survival M9101 43. Stipanovic et al. The female fecundity in the second generation was signiÞcantly lower on M9101.9bA 99.2bB 59. 1964). economical. It has been proved that high-gossypol cotton cultivars are resistant to cotton aphid.001 P 0.9aA 0.9aA 0.05).3 Ϯ 2.1aA 0. 1994.9aB 56.7aA 20.005 0. tabaci MEAM1 cryptic species on the medium (HZ401).9cA 0. tabaci and the results are not consistent (Schuster 1979.0aA 1.9aB 65.001 Ͻ0. tabaci MEAM1 cryptic species has a wide range of host plants. Our present results on B.0bA Ͻ0. tomato fruitworm.8 Ϯ 0.029 P 0. we found that the Þtness of B. with shorter nymphal development time and lower nymphal survival rate.0 Ϯ 3.112 0.476 0. a R ϭ Nt/NtϪ1.001). tabaci MEAM1 cryptic species Indices Male longevity (d) Female longevity (d) No.3aA 100.1 Ϯ 2.7aB 95.2cB 121.3 Ϯ 4.05).0aB 78.2 Ϯ 5.232 HZ401 65.0aA 0.0 Ϯ 0.1aA 0.001 Second-instar M9101 80. and rate of population increase.6 Ϯ 3. and beet armyworm.001 108.6 Ϯ 1. On each cotton cultivar.8 Ϯ 3.7aA 19. tobacco budworms.05 Mean Ϯ SD.: RESPONSE OF B.0 Ϯ 0.005 rate (%) HZ401 38.2aA 20. Because signiÞcantly higher contents of nitrogen. tabaci on nongossypol cultivars. the rate of increase was higher in the second generation than in the Þrst generation (Table 5). 1999.153 Ͻ0.7 Ϯ 3.3 Ϯ 2.2aA 0.7 Ϯ 2. tabaci TO GOSSYPOL 1383 Table 4.7aA 0.3 Ϯ 2.4bA 95.or high (M9101)gossypol cultivars was signiÞcantly lower than that on the low-gossypol cultivar ZMS13.0bA 139.4 Ϯ 14. 2008). which may assist its successful invasion into new ecosystems (Delatte et al.6aA 63.2aA 22. 1964.012 P 0.065 ZMS13 68. 1998).205 survival rate (%) ZMS13 90.2 Ϯ 1. gossypii (Bottger et al.2bA 19. Discussion B. zea (Stipanovic et al. Meng et al.314 (%) ZMS13 97.0 Ϯ 5. compared with that in the Þrst generation (Table 5).0 Ϯ 0.374 survival rate HZ401 84.3 Ϯ 2.05). which was Þrst reported in 1985 in Turkey (Bedford et al.05). Different uppercase letters indicate signiÞcant differences between the two generations on a deÞned cotton cultivar (StudentÕs t-test at P Ͻ 0. It was the highest on ZMS13 and the lowest on M9101. Ozgur and Sekeroglu 1986).0 Ϯ 3.9bA 0.2 Ϯ 1.4bB 98. In each generation of the whiteßy.6 Ϯ 4. Even so. .4aA 100.2 Ϯ 2.0 Ϯ 5. H.7 Ϯ 2. fecundity. and did not differ signiÞcantly on HZ401.9aA 0.3 Ϯ 6.0 Ϯ 3. Different lowercase letters indicate signiÞcant differences among the three cotton cultivars at a deÞned generation (ANOVA: LSD test at P Ͻ 0.3 Ϯ 3.5aA 89.183 0.9 Ϯ 12.008 0. who reported high numbers of B.05 30.103 0. 2006).5 Ϯ 4.2bA 55. Gossypol is one of the most important toxic allelochemicals against insect herbivores in cotton (Bottger et al. the female fecundities of the whiteßy differed significantly among the three cotton cultivars (P Ͻ 0. and free fatty acid were detected in the lowgossypol cultivar ZMS13 in our experiment.019 20.0 Ϯ 3.0aB 0.3 Ϯ 2. Different uppercase letters indicate signiÞcant differences between the two generations on a deÞned cotton cultivar (StudentÕs t-test at P Ͻ 0.018 19. exigua (Wu et al.4bA 117.0aA 18. with HZ401 in between. Effects of cotton cultivar on the fecundity of B.2 Ϯ 18.39 28. There are few studies on the host plant effects of gossypol on B.001 Female M9101 53.June 2013 GUO ET AL.4abA 22.050 Mean Ϯ SD.1 Ϯ 5.3 Ϯ 2. Different lowercase letters indicate signiÞcant differences among the three cotton cultivars at a deÞned generation (ANOVA: LSD test at P Ͻ 0.001 ZMS13 45.9cA 0. Fecundities of Female Adults. Plant secondary metabolites form an important biochemical basis for host plant resistance to insects (Kessler and Baldwin 2002). protein.000 Third-instar HZ401 90.055 0. where NtϪ1 is the number of the initial population and Nt is the number of the future population.0aA 73.9 Ϯ 15.1 Ϯ 3.0aA 0. Table 5.4aA 18.6aA Ͻ0. A.400 percentage (%) HZ401 60.2 Ϯ 26.194 0.
7% on HZ401. the high-gossypol cultivar M9101 may be not only detrimental to sucking pests but also to chewing pests. and R. J. Henneberry. Þeld conditions. Francis. Comparing with that in the Þrst generation. Our results showed that after host plant switch. Insect Biochem. Gao. J. Biol. A. and P. China) for her help during the experiment and Prof. Bemisia argentifolii (Homoptera: Aleyrodidae) populations and relationships to sticky cotton and cotton yields. Res. E. Parajulee. H. J. Invasions 8: 287Ð294.R. F. Brown. 22: 433Ð 449. Naranjo. F.. From our current study. E. Wu et al.S. H. Then. female percentage. References Cited Ascher. Duyck. C. Briddon. P. and S. Our current study provides a proÞle to exemplify the effects of gossypol level on the Þtness of B. De Barro. Showler. when feeding on the high-gossypol cultivar M9101 in three consecutive generations.. Measuring the development. Perkins. Nonconventional insecticidal effects of pesticides available from the neem tree. 3 Perring 2001). Relation of gossypol content of cotton plants to insect resistance. Interactive effects of elevated CO2 and cotton cultivar on tri-trophic interaction of Gossypium hirsutum. F. Physiol. N. 2011. The results derived from this study indicated that B. Becker.. 1993. T. slowdown insecticide resistance development in the whiteßy populations. Arch. 2009. M. L.. De Pauw. Southwest. 56: 1Ð19. L. S. F.1384 JOURNAL OF ECONOMIC ENTOMOLOGY Acknowledgments Vol. R. Environ. P. G. Ge. Entomol. whiteßy cryptic species or populations. 88: 219 Ð229. J. 18: 1Ð10. and rate of population increase in response to gossypol can provide a comprehensive evaluation of plant allelochemicals on the life history of the whiteßy. The adaptation was the slowest on the high-gossypol cultivar M9101. survival rate. 36: 219 Ð227. It indicated an adaptation of the whiteßy to these cotton cultivars during the two generationsÕ rearing. van Lenteren. the rate of increase (R) of the whiteßy was higher in the second generation than in the Þrst generation on each experimental cotton cultivar. Wang. Mol. J. Kang. 106. M. 1994.. Biol. 2006). and T. Invasions 11: 1059 Ð1070. S. and V. 2008. Triboire. Liu. E. Geminivirus transmission and biological characterization of Bemisia tabaci (Gennadius) biotypes from different geographic regions. Drost.. Denmark) for suggestions and linguistic revision of the manuscript. Econ. PLoS ONE 6: e16061.S. T. Jinzhou. W. no. M. Bonato. 45. Bemisia tabaci: a statement of species status. X. S. Harmel. which will reduce pesticide use and residues in food products. J. C. K. . David. 37: 29 Ð37. S. tabaci in these studies might be because of cotton cultivars. Mazzucchelli. R. Khan. P. and shows prosperous application future in cotton pest control practice. (2010) observed significantly longer larval development time. A. Bemisia tabaci: the current situation in the UK and the prospect of developing strategies for eradication using entomopathogens. Bull. Hendrix. and F.W. Ga ´ bor Lo ¨ vei (Aarhus University. Entomol. N. H. By this token. F. Insect Biochem. Gerkens. and National Special Fund for Agricultural Research Grant 201303019. Markham. Rosell. L. Delatte. Cuthbertson. Biol. the insectÕs metabolic adaptation systems will change to cope with the plant secondary substances (Francis et al. Greenberg. Zhao. I. Zhu. R. J. Bottger.. D.. H.. and protect beneÞcial organisms in cotton Þelds (Sharma and Ortiz 2002). tabaci MEAM1 cryptic species over two consecutive generations. Hu. M. Haubruge. the whiteßy adults were collected and released on different cotton cultivars to test the gossypol impacts. lower fecundity. Eyre. The differences of gossypol effects on B. By using the three cotton cultivars (the same as in our current study). A. L. S. 1995. Burke. Reynaud. A. Insect Sci. and A. Such an adaptation was much rapider on the low-gossypol cotton cultivar ZMS13 than that on the medium.or high-gossypol cultivars. J. D. O. K.A. 125: 311Ð325. Bedford. Entomol.9% on M9101. Blackburn. L. B. Brancatini. Northing. Life-history parameters of different biotypes of Bemisia tabaci (Hemiptera: Aleyrodidae) in relation to temperature and host plant: a selective review. Azadirachta indica.L. This work was funded by National Science Foundation of China Grant 30800722. De Barro. Jech.. Dinsdale. 1998. Liu. the colony of B. In the current study. 20: 255Ð271. S. and Propylaea japonica. Akay. Differential invasion success among biotypes: case of Bemisia tabaci. Flint. Bourne. S. Sheehan. tabaci MEAM1 cryptic species was reared on potted tomato plants Goldensun for over 10 generations before the experiment. D..C.J. and relative growth rates of the beet armyworm. P. P. 2005.9% on ZMS13. Boykin. Miller. Y. 2011. We thank Yan-Li Yu (Yangtze University. tabaci MEAM1 cryptic species that fed on the low-gossypol cultivar had a higher Þtness relative to the higher-gossypol cultivars.. Entomol. tabaci MEAM1 cryptic species. 1964. van Roermund. and M. After host plant switch. Lukefahr. Effects of neem-based insecticides on beet armyworm (Lepidoptera: Noctuidae). A. Rev. T. Therefore.J. 2011. the R value in the second generation increased 24. F. G. J. P. Biol. and H. H. Nardi. host plant resistance of a deÞned gossypol cotton cultivar to a deÞned whiteßy cryptic species should be determined before Þeld application. L. Understanding the Þtness parameters involved in the host plantÐ herbivore interaction will aid in exploiting plant allelochemicals in IPM strategies. J. Du. 12: 17Ð23. exigua. P.G. and E. Host plant and biotype density interactions Ð their role in the establishment of the invasive B biotype of Bemisia tabaci. and other factors. and 137. and P. 2006. J. C. T. D. Annu. and B. Proteomics in Myzus persicae: effect of aphid host plant switch. Insect Sci. Aphis gossyppii. fecundity. G.. we conclude that high-gossypol cotton cultivar M9101 can be used in the integrated management program of B. Liu. An extensive Þeld survey combined with a phylogenetic analysis reveals rapid and widespread invasion of two alien whiteßies in China. A. Entomol. Cannon. National Basic Research and Development Program of China Grant 2009CB119200. S. Ann. R. 57: 283Ð285. 2006. Appl. De Barro.
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