Thermal effects on the biological parameters of bean Aphis craccivora (Hemiptera: Aphididae)

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The aim of this study was to investigate the thermal effects on the developmental period, longevity, and fecundity of apterous females of A. craccivora . The experiment was conducted under the effects of four temperature regimes, which included 16°C, 20°C, 24°C, and 28°C, with 65 ± 5% relative humidity (RH) and a photoperiod of long-day 16:8 (L:D) h. Afterwards, the nymphs developed successfully until the adult stage at all temperature regimens. The developmental period of immature A. craccivora ranged from 10.6 days at 16°C to 5.0 days at 28°C. The nymph viability and survival were greater at 24°C than at the other temperatures. However, at a constant temperature of 28°C, the mortality rate was greater than that of the other species at the immature stage of A. craccivora . The lower developmental threshold for cowpea aphids was estimated at 1.77°C and 66.79 degree-days (DD) at the first instar until adulthood. The average longevity of adult females decreased from 22.2 days at 16°C to 10.8 days at 28°C. The net reproduction rate per female was 46.97 at 24°C and 26.93 to 28°C. The largest intrinsic rates of increase ( r m = 0.367) occurred at 28°C, and the smallest occurred at 16°C ( r m = 0.177). It was obvious that temperatures greater than 28°C resulted in good development, increased mortality at the nymphal stage, reduced adult longevity, and diminished fecundity. The optimal growth temperature for A. craccivora was 20°C-24°C. Entomology Behavioral Ecology Developmental Biology Population Biology Adult longevity life table nymphal stage nonlinear function thermal resistance Figures Figure 1 Figure 2 Figure 3 Introduction Aphids (Hemiptera: Aphididae) are cosmopolitan and important pests in agroecosystems and are among the most devastating pests in tropical, subtropical, and temperate regions (Dedryver et al., 2010 ). From this family, the cowpea aphid Aphis craccivora represents one of the most crucial pests affecting the early stages of its host in Africa, Asia, and North America and causing intensive losses in horticultural crops as well as in forestry (Obeng-Ofori., 2007; Ou&edraogo et al., 2018). Parthenogenetic reproduction is the most disastrous damage caused by A. craccivora . The adult winged (alatae) plant causes less damage but is primarily responsible for the infestation of fields through its ability to fly from one place to another. Adults and nymphs feed on the undersurface of young leaves, stem tissues, growing tips, petioles, flowers, and fresh pods by piercing-sucking sap (Togola et al., 2017 ). Throughout the direct and indirect damage caused by A. craccivora , A. craccivora directly feeds by sucking sap and releasing honeydew, which creates fumagine (a sooty mold) on leaves, reducing the plant’s photosynthetic capacity. The bioactive substances injected through saliva interact with plant physiology, disturbing plant growth and development. Indirect damage is the result of virus transmission (Ebert & Cartwright, 1997 ; Blackman & Eastop, 2000 ). Plant damage is increasing because aphids, as virus hosts, are responsible for spreading viral diseases (Aldryhim & Khalil, 1993 ; Smith & Boyko, 2007 ). These include bean necrotic yellows virus, broad bean yellow mosaic virus, and bean leaf roll virus (Weigand & Bishara, 1991 ). The secretion of aphid honeydew reduces photosynthesis by releasing sooty mold on plant leaves (Klingler et al., 2001 ; Smith & Boyko, 2007 ). Among the aforementioned types of aphid damage, abiotic and biotic factors play significant predominant roles in aphid biological life. In particular, abiotic factors affect aphids by modifying their life cycle. Aphids are ectothermic organisms, and abiotic factors strongly affect their development and growth until death. The reported data on the developmental rate and fecundity of cowpea aphids at several temperature regimens revealed different biological variations in Egypt (Hafiz., 2006), Riyadh, Kingdom of Saudi Arabia (Soffan & Aldawood, 2014 ), China (Zhaozhi et al., 2017 ), Korea (Cho et al., 2018 ), and Japon (MOUSA et al., 2019 ). The developmental and fecundity data for all aphid species, from one region to another, should be taken with considerable prudence for different crops because the aphid life table varies with alternating weather conditions. The findings on A. craccivora population parameters can be applied to developing IPM tactics, particularly in monitoring and simplifying the control methods of cowpea aphids in the Eastern Mediterranean region of Turkiye (Kersting et al., 1999 ; Satar et al., 2005 ). The aim of this work was to investigate the life-table parameters of the bean aphid A. craccivora on bean leaves at different temperature regimens under laboratory conditions. Materials and Methods Plant culture : Pinto bean ( Phaseolus vulgaris L.) leaves were collected from the Department of Plant Protection Field Experimental in March 2020 at Adana, Saricam, Turkiye. After collection, the samples were transported to the laboratory and washed below flowing water for 5–10 minutes before use. Every five days, the old leaves were replaced with new leaves for better feeding by the individual insects. Insect culture : Cowpea aphid adults ( A. craccivora) were collected from Acacia trees in the Cukurova University area in Adana, Turkey. On average, 50 adults were reared inside 5 cages on common bean plants ( F. vulgaris L.) at 24 ± 1°C and 65 ± 5% RH with a photoperiod of 16:8 h (L:D). The aphid fabae cages were established on fava bean plants and maintained for five generations prior to the start of the experiment to allow us to recover the net generation from maternal effects reflecting recent rearing conditions. The food A. craccivora was supplied daily to maintain the population stock (Fig. 1 ). Experimental design The experiment was conducted with randomly selected apterous females from stock cage culture and individually transferred to the undersurface of bean leaves on plastic Petri dishes (both 5 cm in diameter). For each temperature, a total of 4 replications of 10 Petri dishes per block with first-instar nymphs were placed inside an incubator. Forty Petri dishes were prepared with a wetted cotton pad (0.5 cm) and placed under the leaves such that the entire surface was covered to avoid drying. After that, 40 newborn aphids were carefully removed with a paintbrush from the master stock to the new Petri dishes. The moisture content of the cotton wool in the Petri dishes was maintained daily, and every 3–5 days, the aphids were transferred to new bean leaf discs. Freshly cultivated leaves were taken from the field and subsequently transported to the Citrus Entomology Laboratory at Cukurova University. The experiments were conducted under four constant temperature regimes (16, 20, 24, and 28 ± 1°C) and 60 ± 5% relative humidity (RH) with a photoperiod of 16:8 (L:D) for 24 hours. For each temperature, the experiment started with 40 first-instar transferred nymphs. Every 24 h, nymphal development was recorded until the adult stage. After the adult period, the number of nymphs and survival of the mother aphids were recorded until the death of all adults of A. craccivora . Statistical analysis The developmental time and reproductive performance of A. craccivora were subjected to analysis of variance (ANOVA). The normality of the data was checked through the Shapiro–Wilk test. Differences in development time, longevity, and reproduction were calculated for each constant temperature. Multiple comparisons were tested using Tukey’s HSD multiple range test (P = 0.05) on significant variables. For each constant temperature, a curve was plotted with the Kaplan‒Meier product limit technique. Population growth rates were computed from the equation of Lotka (Birch, 1948) (Eq. 1). 1 = Σ e- * l* m (1) where x = age in days (including immature stages), r = intrinsic rate of increase, l x = age-specific survival (including immature mortality), m x = age-specific number of female offspring. After "r" was computed for the original data (r all ), differences among the r m values were tested for significance differences by estimating variances through the jackknife method (Meyer et al., 1986). The jackknife pseudo value, r j, was computed for the n samples using the following equation (Eq. 2): r j = n* r all - (n-1) * r i (2) The means of the "n" jackknife pseudo values for each treatment were subjected to analysis of variance. Tukey’s HSD multiple range test was used to compare the mean growth rates at different temperature regimens (P < 0.05). Because low probability levels were used, there was no concern about inflation of experiment wise error rates (Jones, 1984). All of the abovementioned analyses were conducted using the Statgraphics software package version 11.5 (SPSS, Inc., Chicago, IL, USA) (Nie et al. 1975 ) The development rates of the individuals reared at the different temperature levels were calculated via linear regression (y = a ± bx). The means (22°C) of the various temperatures at 16, 20, 24, and 28°C were used in the regression analysis. Afterward, the development threshold (-a/b) and thermal constant (the total effective temperature required to complete a generation, 1/b) of A. craccivora were estimated via linear regression (Campbel et al., 1974). Results The development time of the cowpea aphid A. craccivora significantly decreased with increasing temperature, ranging from 5.0 days at 28°C to 10.6 days at 16°C (F = 81.786; fd = 3; P < 0.05) (Table 1 ). Linear regression analysis was applied to developmental points within the 16°C-28°C range. The temperature range increased linearly with increasing temperature (r (T) = 0.015x – 0.0263; R 2 = 0.7959; F = 245.68; fd = 3; P < 0.05) (Fig. 1 ). The lower developmental threshold (LT) and thermal constant (K) of the A. craccivora nymphal stage were estimated to be 1.77°C, and 66.79 degree days (DD) were required for the first instar to become adults (Table 3 ). The longevity of the plants was significantly greater at 16°C (F = 17.858; fd = 3; P < 0.05) (Table 2 ). compared to any other temperature regime tested. The constant temperature for the highest number of offspring days occurred at 24°C (F = 2.74; fd = 3; P < 0.05) (Table 3 ). The highest average value of fecundity per reproductive day occurred at 24°C (F = 1.811; fd = 3; P < 0.05), and the lowest was at 16°C (Table 2 ). Table 1 Development times (days ± SE) of Aphis craccivora on bean ( Phaseolus vulgaris . L) at five constant temperatures, 65 ± 5% RH, and a photoperiod of 16:00 (L: D) h. Temperature (°C) n I. Nymph period II. Nymph period III. Nymph period IV. Nymph period Total dev. 16°C 40 2.7 ± 0.14a 2.6 ± 0.16a 2.4 ± 0.31a 3.2 ± 0.28a 10.6 ± 0.42a 20°C 40 1.4 ± 0.08b 1.6 ± 0.12b 1.1 ± 0.10 c 1.7 ± 0.17b 6.0 ± 0.30b 24°C 40 1.0 ± 0.00c 1.1 ± 0.04c 1.8 ± 0.07ab 1.4 ± 0.10b 5.2 ± 0.16b 28°C 40 1.1 ± 0.05c 1.0 ± 0.06c 1.5 ± 0.12 bc 1.3 ± 0.12b 5.0 ± 0.17b Significant differences between means (*P < 0.05 and **P < 0.01) are expressed by different letters (a–b). The letters compare values in the same column. Table 2 Preoviposition, oviposition, postoviposition, longevity, life span, and number of adult female Aphis craccivora individuals on bean (mean ± SE) Temp (°C) Pre-Oviposition (day) Oviposition (day) Post-Oviposition (day) Longevity (day) Life span (day) Offspring (Aphid) 16 1.17 ± 0.2a 8.95 ± 1.5a 0.25 ± 0.1 22.2 ± 1.71a 11.6 ± 1.75a 32.1 ± 5.4 20 0.85 ± 0.1ab 6.77 ± 0.9ab 0.35 ± 0.4 14.1 ± 1.29b 8.1 ± 1.20ab 40.8 ± 6.2 24 0.45 ± 0.1b 6.32 ± 0.4ab 0.42 ± 0.1 12.4 ± 0.49b 7.2 ± 0.50b 45.2 ± 3.4 28 0.55 ± 0.1b 5.07 ± 0.8b 0.45 ± 0.1 10.8 ± 0.89b 5.8 ± 0.86b 26.9 ± 4.5 Significant differences between means ( * P < 0.05 and ** P < 0.01) are expressed by different letters (a–c). The letters compare values in the same column. Table 3 Offspring/day, death ratio (%), wingless adults (n), wing adults (n), generation time (T0), net reproduction (Ro), and intrinsic rate of increase (rm) of Aphis craccivora on bean leaf discs at five temperature levels Temp (°C) N Offspring reproductıon day(mean ± SE) Death ratio (%) Wingless adults(N) Wing Adult(N) (T 0 ) (R 0 ) (r m ) 16 40 3.6 ± 1.77 20 36 4 22.287 32.175 0.177 20 40 4.2 ± 0.47 20 40 3 13.294 40.850 0.321 24 40 6.5 ± 0.45 5 38 4 13.174 46.975 0.352 28 40 4.1 ± 0.45 28 35 5 10.191 26.925 0.367 Aphis craccivora (a) fourth nymphal stage, (b) aphid exoskeleton, skin or exuvia stage, (c) adult stage, and (d) first nymphal stage. The highest % mortality rate was observed at 28°C, possibly because of the vulnerability to high temperature at the first nymphal stage (Table 3 ). The survival rate of A. craccivora adults sharply diminished after the peak of nymph production at higher temperatures. More wingless adults were observed at 20°C. However, at 28°C, there were more winged or alate plants (Table 3 ). According to the biological development of aphid species at warm temperatures, the possibility of developing winged individuals is greater. At 16°C and 20°C, the post oviposition period was relatively long compared to that at the other temperatures. (Fig. 3 ) (Table 2 ). The number of reproductive offspring per period of day varied between 3.6 days (16°C) and 6.5 days (24°C) (Table 3 ). Augmenting the temperature resulted in shorter generation times ( T o ) of A. craccivora , with 22.3 days at 16°C and 10.2 at 28°C (Table 3 ). The net reproduction rate ( R o ) was highest at 24°C (64.97 aphids/aphid) and lowest at 28°C (26.93 aphids/aphid) (Table 3 ). A greater population of A. craccivora resulted in a greater per capita rate of population growth, as indicated by the intrinsic rate of increase at 28°C (0.367 aphids/aphid per day) compared to that at 16°C (0.177 aphids/aphid) (Table 3 ). The developmental rate (r) of A. craccivora at four constant temperatures, all of which were alternated (variation cycle), was fitted with a linear regression equation, and the developmental rate ranged between 16°C and 28°C (Table 4 ). The mean alternating temperatures were used to fit the linear regression equation. The development time of A. craccivora increased linearly with increasing temperature. The development rates of A. craccivora at four temperature regimens were fitted to the linear regression equation y = a ± bx (Table 4 ). The outcomes of the regression model were fitted separately for the data obtained from female A. craccivora at the developmental rate. The equation for female development was calculated as follows: Y = 0.015 × -0.0263 (R2 = 0.97; P ≤ 0.05) from the first instar to the developmental stage and adult stage (Fig. 2 ): through these equations, the development thresholds and thermal constants were calculated. Table 4 Regression equations and parameters of development period rates of Aphis craccivora on bean leaves under different constant temperatures Parameters Nymph I Nymph II Nymph III Nymph IV Adult Equation of regression y = -0.13x + 4.4 R² =0.730 y= -0.13x + 4.49 R² = 0.873 y = -0.05x + 2.8 R² = 0.222 y = -0.15x + 5.2 R² = 0.769 y = -0.44x + 16.38 R² = 0.743 Thermal constant (1/a) (°C day) 7.69 7.54 20 6.67 2.27 Development threshold (b/a) (°C) 260.08 225.50 1120 231.22 84.51 Table 5 ANOVA of several biological parameters of Aphis craccivora at five temperature regimens on the bean. A P value less than 0.05 indicated that there were significant differences between parameters in this study. As we can see only for offspring per day, postoviposition, and lifespan, there was no significant difference because the p value was > 0.05. Biological parameters Between Groups Sum of Squares df Mean Square F Sig. Lifespan 726.069 3 242.023 4.340 0.006‘**’ Offspring 8192.725 3 2730.908 2.744 0.045 ʿ*ʾ Longevity 3043.850 3 1014.617 17.858 0.000*** N1 79.269 3 26.423 80.311 0.000*** N2 69.525 3 23.175 44.938 0.000*** N3 35.469 3 11.823 9.221 0.000*** N4 93.869 3 31.290 22.590 0.000*** Total-development 824.269 3 274.756 81.786 0.000*** Preoviposition 12.819 3 4.273 5.024 0.002‘**’ Oviposition 312.919 3 104.306 2.519 0.060‘*’ Postoviposition 85.119 3 28.373 10.715 0.000*** Offspring/day 205.626 3 68.542 1.811 0.147‘.’ Discussion In the ecosystem, insects are not subjected to constant or alternating temperatures. However, laboratory conditions can provide valuable insight into the population dynamics of aphids. The findings reported here clearly show the effects of temperature on the development time, death rate, longevity, and fecundity of A. craccivora . Aphid species are small ectothermic insects that are strongly related to temperature. Recall that temperature influences multiple aspects of insect biology, such as the metabolic system and developmental rate, as well as the timing and level of insect activities. Temperatures beyond the thresholds of development for a given insect can slow growth, and extreme temperatures can kill members of the insect population. It is logical that temperature fluctuations can cause changes in insect biology. In this study, we have discussed how temperature alternations affect insect biology. Several biological parameters of Aphis craccivora were also evaluated. The maximum temperature for the development of A. craccivora was 28°C (mean = 5.0 days; F = 81.78; P = 0.05), but other studies have reported a temperature of 30°C (Cho et al., 2018 ; Kuo, M. H. & Chen, 2004) and a temperature of 29.4°C (Berberet et al., 2009 ; Girão et al., 2019 ) on bean plants. These results correlate with the degree to which ectothermic animals that develop under warm conditions tend to grow faster and mature earlier. On the other hand, these animals are slower in maturation than are similar animals that develop under cool conditions. The preoviposition period, during which the temperature is held constant and gradually decreases, delays the occurrence of fecundity. The temperature increases from 16°C to 24°C, similar to the 16°C and 25°C reported by (Cho et al., 2018 ; Girão et al., 2019 ). The number of oviposition days strongly depends on the area in which the insect is evaluated by using food and weather resources. Normally, at both low- and high-tolerance temperatures, insects can take a long time to reproduce, for example, from 16°C to 28°C (mean = 8.95, 5.07; F = 2.519; P = 0.05). The same tendency at 15, 20, and 25°C was reported by (Cho and et al., 2018 ) and at 18, 22, 25, and 28°C. (Girão et al., 2019 ). The postoviposition period was 16°C (mean day = 2.05; F = 2.519; P = 0.05) if it was below the lowest relative temperature for good reproduction; during this time, the duration after the offspring period became longer than the adaptive temperatures reported at 18, 22, 25, and 28°C (Girão, et al., 2019 ) (mean days = 1.2, 0.8, 0.8, and 0.2, respectively). Longevity increases inversely with decreasing temperature, as we can see at 16°C (22.2 days) and 28°C (10.8 days); however, in the literature, approximately the same constant temperatures were reported at 18°C and 28°C by (Girão et al., 2019 ). However, there were differences between the results presented in this research within the life span of A. craccivora . The temperature regimes exhibited significant differences from the highest to lowest temperature at 28°C and 16°C (mean/day) (5.8 and 11.6, F = 4.340; P = 0.05, respectively). The mean values between the life span temperatures were similar: at 28°C, the lowest occurred at 16°C, and the highest occurred at 28°C. These findings revealed that there were no significant differences. These results indicate that these temperatures are amenable to survival for aphid species. (Table 2 ). In general, these observations established the inverse proportional relationship between oviposition periods and immature stages produced per female per day in the temperature range of 16 to 28°C, in contrast to the results presented by(Berberet et al., 2009 ; Girão et al., 2019 ). According to these results, there are many possible adaptations between several studied aphid populations. For example, the variations observed in thermal constant development based on the suitable temperature for black aphids on the bean might be explained in part by the functional theory of life, which relates to the highest value for a development threshold and the smallest value for total thermal requirement, as expected, for species more adapted to tropical regions and temperate regions (Trudgill & Perry, 1994 ; Brown et al., 1995 ). Declarations Conflict of interest The authors declare that they have no conflicts of interest. Ethical approval This article does not contain any studies with human participants performed by any of the authors. Acknowledgments We would like to thank the team management of the Project Development and Coordination Unit. This research was part of the Master’s degree thesis of Rochelyn DONA and was funded by the Project Development and Coordination Unit of Cukurova University (Grant Code: FYL-2020-12688). 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Ann Appl Biol no 3:521–532 Weigand S, Bishara SI (1991) Status of insect pests of faba bean in the Mediterranean region and methods of control. Serie A: Seminaires Mediterraneens Zhaozhi L, Likai F, Guizhen G, Ling-Ling G, Han P, Sharma S, Zalucki MP (2017) Differences in the high-temperature tolerance of Aphis craccivora (Hemiptera: Aphididae) on cotton and soybean: implications for ecological niche switching among hosts. Appl Entomol Zool 52:9–18 Additional Declarations The authors declare no competing interests. Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-4960855","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":343933169,"identity":"4a817915-1879-4a93-9258-da6c47d62bd7","order_by":0,"name":"Rochelyn Dona","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA+klEQVRIiWNgGAWjYDCCAxAqgYGdsYHhQ4UNkM3YeIA4LcyMDYwzzqSBtDQQq4WBgZm37TCyIHbAd7z34OeKCrs8fmbmtocz287brW0/DLSlxiYalxbJM+eSJc+cSS6WbGZsN/hw7nbytjOJQC3H0nIbcGgxuJFjINnYdiBxw2HGNskZZbeTzQ4AtTA2HManxfgnSMt+oBZpHrZzyWbnHxLUYgaxhRmkpe2AndkNArYA/WFm2XAmOXEG2GFnkhPMbgBtScDjF77jPcY3GyrsEvvb259JfKiwszc7n/7wwYcaG5xaMEAiWGUCscpBwJ4UxaNgFIyCUTAyAAD4BmtKMa2zQwAAAABJRU5ErkJggg==","orcid":"https://orcid.org/0000-0002-4868-8356","institution":"Cukurova University","correspondingAuthor":true,"prefix":"","firstName":"Rochelyn","middleName":"","lastName":"Dona","suffix":""},{"id":343933170,"identity":"8e070aa1-28f7-4fcc-a0a0-4bbff4e13b02","order_by":1,"name":"Serdar Satar","email":"","orcid":"","institution":"Cukurova University","correspondingAuthor":false,"prefix":"","firstName":"Serdar","middleName":"","lastName":"Satar","suffix":""}],"badges":[],"createdAt":"2024-08-23 02:09:56","currentVersionCode":1,"declarations":{"humanSubjects":true,"vertebrateSubjects":true,"conflictsOfInterestStatement":false,"humanSubjectEthicalGuidelines":true,"humanSubjectConsent":true,"humanSubjectClinicalTrial":false,"humanSubjectCaseReport":false,"vertebrateSubjectEthicalGuidelines":true},"doi":"10.21203/rs.3.rs-4960855/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4960855/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":63254514,"identity":"27231dfa-9b86-4e35-8d32-576a6c778738","added_by":"auto","created_at":"2024-08-26 07:53:13","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":310532,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eAphis craccivora\u003c/em\u003e (a) fourth nymphal stage; (b) aphid exoskeleton, skin or exuvia stage; (c) adult stage; and (d) first nymphal stage\u003c/p\u003e","description":"","filename":"Fig.1.png","url":"https://assets-eu.researchsquare.com/files/rs-4960855/v1/59628567ed9a52c296c1dd26.png"},{"id":63255263,"identity":"c4c7730e-4740-463a-9868-3931f44442ae","added_by":"auto","created_at":"2024-08-26 08:01:13","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":32378,"visible":true,"origin":"","legend":"\u003cp\u003eExplaining the developmental severity of \u003cem\u003eAphis craccivora\u003c/em\u003e on four (4) temperature regimens.\u003c/p\u003e","description":"","filename":"Fig.2.png","url":"https://assets-eu.researchsquare.com/files/rs-4960855/v1/ae93f52f0daefbf0b236ee76.png"},{"id":63254512,"identity":"3b5a97e0-86b6-42ae-ad9d-d2552558effb","added_by":"auto","created_at":"2024-08-26 07:53:13","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":81450,"visible":true,"origin":"","legend":"\u003cp\u003eAge-specific survival rate (l\u003csub\u003ex\u003c/sub\u003e) and fecundity (m\u003csub\u003ex\u003c/sub\u003e) of\u003cem\u003e Aphis craccivora \u003c/em\u003ereared on bean leaves (\u003cem\u003ePhaseolus vulgaris\u003c/em\u003e L.) at four constant temperatures\u003c/p\u003e","description":"","filename":"Fig.3.png","url":"https://assets-eu.researchsquare.com/files/rs-4960855/v1/6c7b87010685edd728db91d1.png"},{"id":63255772,"identity":"b33804fb-a2c9-4f3c-9575-9c53733f6fc3","added_by":"auto","created_at":"2024-08-26 08:09:13","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":988657,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4960855/v1/e24021c1-1f36-4754-a22c-da560779d185.pdf"}],"financialInterests":"The authors declare no competing interests.","formattedTitle":"\u003cp\u003e\u003cstrong\u003eThermal effects on the biological parameters of bean \u003c/strong\u003e\u003cem\u003e\u003cstrong\u003eAphis craccivora\u003c/strong\u003e\u003c/em\u003e\u003cstrong\u003e (Hemiptera: Aphididae)\u003c/strong\u003e\u003c/p\u003e","fulltext":[{"header":"Introduction","content":"\u003cp\u003eAphids (Hemiptera: Aphididae) are cosmopolitan and important pests in agroecosystems and are among the most devastating pests in tropical, subtropical, and temperate regions (Dedryver et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). From this family, the cowpea aphid \u003cem\u003eAphis craccivora\u003c/em\u003e represents one of the most crucial pests affecting the early stages of its host in Africa, Asia, and North America and causing intensive losses in horticultural crops as well as in forestry (Obeng-Ofori., 2007; Ou\u0026amp;edraogo et al., 2018). Parthenogenetic reproduction is the most disastrous damage caused by \u003cem\u003eA. craccivora\u003c/em\u003e. The adult winged (alatae) plant causes less damage but is primarily responsible for the infestation of fields through its ability to fly from one place to another. Adults and nymphs feed on the undersurface of young leaves, stem tissues, growing tips, petioles, flowers, and fresh pods by piercing-sucking sap (Togola et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Throughout \u003cem\u003ethe\u003c/em\u003e direct and indirect damage caused by \u003cem\u003eA. craccivora\u003c/em\u003e, \u003cem\u003eA. craccivora\u003c/em\u003e directly feeds by sucking sap and releasing honeydew, which creates fumagine (a sooty mold) on leaves, reducing the plant\u0026rsquo;s photosynthetic capacity. The bioactive substances injected through saliva interact with plant physiology, disturbing plant growth and development. Indirect damage is the result of virus transmission (Ebert \u0026amp; Cartwright, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Blackman \u0026amp; Eastop, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). Plant damage is increasing because aphids, as virus hosts, are responsible for spreading viral diseases (Aldryhim \u0026amp; Khalil, \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Smith \u0026amp; Boyko, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). These include bean necrotic yellows virus, broad bean yellow mosaic virus, and bean leaf roll virus (Weigand \u0026amp; Bishara, \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e1991\u003c/span\u003e). The secretion of aphid honeydew reduces photosynthesis by releasing sooty mold on plant leaves (Klingler et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Smith \u0026amp; Boyko, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2007\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAmong the aforementioned types of aphid damage, abiotic and biotic factors play significant predominant roles in aphid biological life. In particular, abiotic factors affect aphids by modifying their life cycle. Aphids are ectothermic organisms, and abiotic factors strongly affect their development and growth until death. The reported data on the developmental rate and fecundity of cowpea aphids at several temperature regimens revealed different biological variations in Egypt (Hafiz., 2006), Riyadh, Kingdom of Saudi Arabia (Soffan \u0026amp; Aldawood, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), China (Zhaozhi et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), Korea (Cho et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), and Japon (MOUSA et al., \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The developmental and fecundity data for all aphid species, from one region to another, should be taken with considerable prudence for different crops because the aphid life table varies with alternating weather conditions. The findings on \u003cem\u003eA. craccivora\u003c/em\u003e population parameters can be applied to developing IPM tactics, particularly in monitoring and simplifying the control methods of cowpea aphids in the Eastern Mediterranean region of Turkiye (Kersting et al., \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Satar et al., \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). The aim of this work was to investigate the life-table parameters of the bean aphid \u003cem\u003eA. craccivora\u003c/em\u003e on bean leaves at different temperature regimens under laboratory conditions.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cp\u003e \u003cb\u003ePlant culture\u003c/b\u003e: Pinto bean (\u003cem\u003ePhaseolus vulgaris\u003c/em\u003e L.) leaves were collected from the Department of Plant Protection Field Experimental in March 2020 at Adana, Saricam, Turkiye. After collection, the samples were transported to the \u003cb\u003elaboratory\u003c/b\u003e and washed below flowing water for 5\u0026ndash;10 minutes before use. Every five days, the old leaves were replaced with new leaves for better feeding by the individual insects.\u003c/p\u003e \u003cp\u003e \u003cb\u003eInsect culture\u003c/b\u003e: Cowpea aphid adults (\u003cem\u003eA. craccivora)\u003c/em\u003e were collected from Acacia trees in the Cukurova University area in Adana, Turkey. On average, 50 adults were reared inside 5 cages on common bean plants (\u003cem\u003eF. vulgaris\u003c/em\u003e L.) at 24\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u0026deg;C and 65\u0026thinsp;\u0026plusmn;\u0026thinsp;5% RH with a photoperiod of 16:8 h (L:D). The aphid fabae cages were established on fava bean plants and maintained for five generations prior to the start of the experiment to allow us to recover the net generation from maternal effects reflecting recent rearing conditions. The food \u003cem\u003eA. craccivora\u003c/em\u003e was supplied daily to maintain the population stock (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cb\u003eExperimental design\u003c/b\u003e \u003c/p\u003e \u003cp\u003eThe experiment was conducted with randomly selected apterous females from stock cage culture and individually transferred to the undersurface of bean leaves on plastic Petri dishes (both 5 cm in diameter). For each temperature, a total of 4 replications of 10 Petri dishes per block with first-instar nymphs were placed inside an incubator. Forty Petri dishes were prepared with a wetted cotton pad (0.5 cm) and placed under the leaves such that the entire surface was covered to avoid drying. After that, 40 newborn aphids were carefully removed with a paintbrush from the master stock to the new Petri dishes. The moisture content of the cotton wool in the Petri dishes was maintained daily, and every 3\u0026ndash;5 days, the aphids were transferred to new bean leaf discs. Freshly cultivated leaves were taken from the field and subsequently transported to the Citrus Entomology Laboratory at Cukurova University.\u003c/p\u003e \u003cp\u003eThe experiments were conducted under four constant temperature regimes (16, 20, 24, and 28\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u0026deg;C) and 60\u0026thinsp;\u0026plusmn;\u0026thinsp;5% relative humidity (RH) with a photoperiod of 16:8 (L:D) for 24 hours. For each temperature, the experiment started with 40 first-instar transferred nymphs. Every 24 h, nymphal development was recorded until the adult stage. After the adult period, the number of nymphs and survival of the mother aphids were recorded until the death of all adults of \u003cem\u003eA. craccivora\u003c/em\u003e.\u003c/p\u003e \u003cp\u003e \u003cb\u003eStatistical analysis\u003c/b\u003e \u003c/p\u003e \u003cp\u003eThe developmental time and reproductive performance of \u003cem\u003eA. craccivora\u003c/em\u003e were subjected to analysis of variance (ANOVA). The normality of the data was checked through the Shapiro\u0026ndash;Wilk test. Differences in development time, longevity, and reproduction were calculated for each constant temperature. Multiple comparisons were tested using Tukey\u0026rsquo;s HSD multiple range test (P\u0026thinsp;=\u0026thinsp;0.05) on significant variables. For each constant temperature, a curve was plotted with the Kaplan‒Meier product limit technique. Population growth rates were computed from the equation of Lotka (Birch, 1948) (Eq.\u0026nbsp;1).\u003c/p\u003e\n\u003cp\u003e1 = Σ e- * l* m (1)\u003c/p\u003e\n\u003cp\u003ewhere x\u0026thinsp;=\u0026thinsp;age in days (including immature stages), r\u0026thinsp;=\u0026thinsp;intrinsic rate of increase,\u003c/p\u003e \u003cp\u003el\u003csub\u003ex\u003c/sub\u003e= age-specific survival (including immature mortality), m\u003csub\u003ex\u003c/sub\u003e = age-specific number of female offspring. After \"r\" was computed for the original data (r\u003csub\u003eall\u003c/sub\u003e), differences among the r\u003csub\u003em\u003c/sub\u003e values were tested for significance differences by estimating variances through the jackknife method (Meyer et al., 1986). The jackknife pseudo value, r\u003csub\u003ej,\u003c/sub\u003e was computed for the \u003cem\u003en\u003c/em\u003e samples using the following equation (Eq.\u0026nbsp;2):\u003c/p\u003e \u003cp\u003er\u003csub\u003ej\u003c/sub\u003e = n* r\u003csub\u003eall\u003c/sub\u003e - (n-1) * r\u003csub\u003ei\u003c/sub\u003e (2)\u003c/p\u003e \u003cp\u003eThe means of the \"n\" jackknife pseudo values for each treatment were subjected to analysis of variance. Tukey\u0026rsquo;s HSD multiple range test was used to compare the mean growth rates at different temperature regimens (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05). Because low probability levels were used, there was no concern about inflation of experiment wise error rates (Jones, 1984). All of the abovementioned analyses were conducted using the Statgraphics software package version 11.5 (SPSS, Inc., Chicago, IL, USA) (Nie et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1975\u003c/span\u003e)\u003c/p\u003e \u003cp\u003eThe development rates of the individuals reared at the different temperature levels were calculated via linear regression (y\u0026thinsp;=\u0026thinsp;a\u0026thinsp;\u0026plusmn;\u0026thinsp;bx). The means (22\u0026deg;C) of the various temperatures at 16, 20, 24, and 28\u0026deg;C were used in the regression analysis. Afterward, the development threshold (-a/b) and thermal constant (the total effective temperature required to complete a generation, 1/b) of \u003cem\u003eA. craccivora\u003c/em\u003e were estimated via linear regression (Campbel et al., 1974).\u003c/p\u003e "},{"header":"Results","content":"\u003cp\u003eThe development time of the cowpea aphid \u003cem\u003eA. craccivora\u003c/em\u003e significantly decreased with increasing temperature, ranging from 5.0 days at 28\u0026deg;C to 10.6 days at 16\u0026deg;C (F\u0026thinsp;=\u0026thinsp;81.786; fd\u0026thinsp;=\u0026thinsp;3; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Linear regression analysis was applied to developmental points within the 16\u0026deg;C-28\u0026deg;C range. The temperature range increased linearly with increasing temperature (r \u003csub\u003e(T)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;0.015x \u0026ndash; 0.0263; R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.7959; F\u0026thinsp;=\u0026thinsp;245.68; fd\u0026thinsp;=\u0026thinsp;3; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The lower developmental threshold (LT) and thermal constant (K) of the \u003cem\u003eA. craccivora\u003c/em\u003e nymphal stage were estimated to be 1.77\u0026deg;C, and 66.79 degree days (DD) were required for the first instar to become adults (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The longevity of the plants was significantly greater at 16\u0026deg;C (F\u0026thinsp;=\u0026thinsp;17.858; fd\u0026thinsp;=\u0026thinsp;3; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). compared to any other temperature regime tested. The constant temperature for the highest number of offspring days occurred at 24\u0026deg;C (F\u0026thinsp;=\u0026thinsp;2.74; fd\u0026thinsp;=\u0026thinsp;3; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The highest average value of fecundity per reproductive day occurred at 24\u0026deg;C (F\u0026thinsp;=\u0026thinsp;1.811; fd\u0026thinsp;=\u0026thinsp;3; \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05), and the lowest was at 16\u0026deg;C (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eDevelopment times (days\u0026thinsp;\u0026plusmn;\u0026thinsp;SE) of \u003cem\u003eAphis craccivora\u003c/em\u003e on bean (\u003cem\u003ePhaseolus vulgaris\u003c/em\u003e. L) at five constant temperatures, 65\u0026thinsp;\u0026plusmn;\u0026thinsp;5% RH, and a photoperiod of 16:00 (L: D) h.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTemperature\u003c/p\u003e \u003cp\u003e(\u0026deg;C)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003en\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eI. Nymph period\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eII. Nymph period\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eIII. Nymph period\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eIV. Nymph period\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eTotal dev.\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e16\u0026deg;C\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.14a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.16a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.31a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.28a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e10.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.42a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e20\u0026deg;C\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.10 c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.17b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e6.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.30b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e24\u0026deg;C\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.10b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e5.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.16b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e28\u0026deg;C\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12 bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e5.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.17b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"7\"\u003eSignificant differences between means (*P\u0026thinsp;\u0026lt;\u0026thinsp;0.05 and **P\u0026thinsp;\u0026lt;\u0026thinsp;0.01) are expressed by different letters (a\u0026ndash;b). The letters compare values in the same column.\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003ePreoviposition, oviposition, postoviposition, longevity, life span, and number of adult female \u003cem\u003eAphis craccivora\u003c/em\u003e individuals on bean (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SE)\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTemp\u003c/p\u003e \u003cp\u003e(\u0026deg;C)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePre-Oviposition\u003c/p\u003e \u003cp\u003e(day)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eOviposition\u003c/p\u003e \u003cp\u003e(day)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePost-Oviposition\u003c/p\u003e \u003cp\u003e(day)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eLongevity (day)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eLife span (day)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eOffspring (Aphid)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e16\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.17\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e8.95\u0026thinsp;\u0026plusmn;\u0026thinsp;1.5a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.25\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e22.2\u0026thinsp;\u0026plusmn;\u0026thinsp;1.71a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e11.6\u0026thinsp;\u0026plusmn;\u0026thinsp;1.75a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e32.1\u0026thinsp;\u0026plusmn;\u0026thinsp;5.4\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e20\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.85\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e6.77\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.35\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e14.1\u0026thinsp;\u0026plusmn;\u0026thinsp;1.29b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e8.1\u0026thinsp;\u0026plusmn;\u0026thinsp;1.20ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e40.8\u0026thinsp;\u0026plusmn;\u0026thinsp;6.2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e24\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.45\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e6.32\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.42\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e12.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.49b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e7.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.50b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e45.2\u0026thinsp;\u0026plusmn;\u0026thinsp;3.4\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e28\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.55\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5.07\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.45\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e10.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.89b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e5.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.86b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e26.9\u0026thinsp;\u0026plusmn;\u0026thinsp;4.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"7\"\u003eSignificant differences between means (\u003csup\u003e*\u003c/sup\u003e\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05 and \u003csup\u003e**\u003c/sup\u003e\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.01) are expressed by different letters (a\u0026ndash;c). The letters compare values in the same column.\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eOffspring/day, death ratio (%), wingless adults (n), wing adults (n), generation time (T0), net reproduction (Ro), and intrinsic rate of increase (rm) of \u003cem\u003eAphis craccivora\u003c/em\u003e on bean leaf discs at five temperature levels\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"9\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTemp\u003c/p\u003e \u003cp\u003e(\u0026deg;C)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eOffspring reproductıon day(mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SE)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eDeath ratio (%)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eWingless adults(N)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eWing Adult(N)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003e(T\u003csub\u003e0\u003c/sub\u003e)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003e(R\u003csub\u003e0\u003c/sub\u003e)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003e(r\u003csub\u003em\u003c/sub\u003e)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e16\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e3.6\u0026thinsp;\u0026plusmn;\u0026thinsp;1.77\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e22.287\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e32.175\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.177\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e20\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e4.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.47\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e13.294\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e40.850\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.321\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e24\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e6.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e13.174\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e46.975\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.352\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003e28\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e4.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e10.191\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e26.925\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0.367\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"9\"\u003e\u003cem\u003eAphis craccivora\u003c/em\u003e (a) fourth nymphal stage, (b) aphid exoskeleton, skin or exuvia stage, (c) adult stage, and (d) first nymphal stage.\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eThe highest % mortality rate was observed at 28\u0026deg;C, possibly because of the vulnerability to high temperature at the first nymphal stage (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The survival rate of \u003cem\u003eA. craccivora\u003c/em\u003e adults sharply diminished after the peak of nymph production at higher temperatures. More wingless adults were observed at 20\u0026deg;C. However, at 28\u0026deg;C, there were more winged or alate plants (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). According to the biological development of aphid species at warm temperatures, the possibility of developing winged individuals is greater. At 16\u0026deg;C and 20\u0026deg;C, the post oviposition period was relatively long compared to that at the other temperatures. (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e) (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The number of reproductive offspring per period of day varied between 3.6 days (16\u0026deg;C) and 6.5 days (24\u0026deg;C) (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Augmenting the temperature resulted in shorter generation times (\u003cem\u003eT\u003c/em\u003e\u003csub\u003e\u003cem\u003eo\u003c/em\u003e\u003c/sub\u003e) of \u003cem\u003eA. craccivora\u003c/em\u003e, with 22.3 days at 16\u0026deg;C and 10.2 at 28\u0026deg;C (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The net reproduction rate (\u003cem\u003eR\u003c/em\u003e\u003csub\u003e\u003cem\u003eo\u003c/em\u003e\u003c/sub\u003e) was highest at 24\u0026deg;C (64.97 aphids/aphid) and lowest at 28\u0026deg;C (26.93 aphids/aphid) (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). A greater population of \u003cem\u003eA. craccivora\u003c/em\u003e resulted in a greater per capita rate of population growth, as indicated by the intrinsic rate of increase at 28\u0026deg;C (0.367 aphids/aphid per day) compared to that at 16\u0026deg;C (0.177 aphids/aphid) (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe developmental rate (r) of \u003cem\u003eA. craccivora\u003c/em\u003e at four constant temperatures, all of which were alternated (variation cycle), was fitted with a linear regression equation, and the developmental rate ranged between 16\u0026deg;C and 28\u0026deg;C (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The mean alternating temperatures were used to fit the linear regression equation. The development time of \u003cem\u003eA. craccivora\u003c/em\u003e increased linearly with increasing temperature. The development rates of \u003cem\u003eA. craccivora\u003c/em\u003e at four temperature regimens were fitted to the linear regression equation y\u0026thinsp;=\u0026thinsp;a\u0026thinsp;\u0026plusmn;\u0026thinsp;bx (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The outcomes of the regression model were fitted separately for the data obtained from female \u003cem\u003eA. craccivora at the\u003c/em\u003e developmental rate. The equation for female development was calculated as follows: Y\u0026thinsp;=\u0026thinsp;0.015 \u0026times; -0.0263 (R2\u0026thinsp;=\u0026thinsp;0.97; P\u0026thinsp;\u0026le;\u0026thinsp;0.05) from the first instar to the developmental stage and adult stage (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e): through these equations, the development thresholds and thermal constants were calculated.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eRegression equations and parameters of development period rates of \u003cem\u003eAphis craccivora\u003c/em\u003e on bean leaves under different constant temperatures\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eParameters\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNymph I\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNymph II\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eNymph III\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eNymph IV\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAdult\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eEquation of regression\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ey = -0.13x\u0026thinsp;+\u0026thinsp;4.4\u003c/p\u003e \u003cp\u003eR\u0026sup2; =0.730\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ey= -0.13x\u0026thinsp;+\u0026thinsp;4.49\u003c/p\u003e \u003cp\u003eR\u0026sup2; = 0.873\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ey = -0.05x\u0026thinsp;+\u0026thinsp;2.8\u003c/p\u003e \u003cp\u003eR\u0026sup2; = 0.222\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ey = -0.15x\u0026thinsp;+\u0026thinsp;5.2\u003c/p\u003e \u003cp\u003eR\u0026sup2; = 0.769\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ey = -0.44x\u0026thinsp;+\u0026thinsp;16.38\u003c/p\u003e \u003cp\u003eR\u0026sup2; = 0.743\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eThermal constant (1/a) (\u0026deg;C day)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7.69\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e7.54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e6.67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2.27\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eDevelopment threshold (b/a) (\u0026deg;C)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e260.08\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e225.50\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1120\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e231.22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e84.51\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab5\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 5\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eANOVA of several biological parameters of \u003cem\u003eAphis craccivora\u003c/em\u003e at five temperature regimens on the bean. A P value less than 0.05 indicated that there were significant differences between parameters in this study. As we can see only for offspring per day, postoviposition, and lifespan, there was no significant difference because the p value was \u0026gt;\u0026thinsp;0.05.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBiological parameters\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"12\" rowspan=\"13\"\u003e \u003cp\u003eBetween Groups\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSum of Squares\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003edf\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMean Square\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eF\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSig.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eLifespan\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e726.069\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e242.023\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4.340\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.006\u0026lsquo;**\u0026rsquo;\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eOffspring\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e8192.725\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2730.908\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2.744\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.045 ʿ*ʾ\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eLongevity\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3043.850\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1014.617\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e17.858\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.000***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eN1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e79.269\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e26.423\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e80.311\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.000***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eN2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e69.525\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e23.175\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e44.938\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.000***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eN3\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e35.469\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e11.823\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e9.221\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.000***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eN4\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e93.869\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e31.290\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e22.590\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.000***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eTotal-development\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e824.269\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e274.756\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e81.786\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.000***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ePreoviposition\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e12.819\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4.273\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e5.024\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.002\u0026lsquo;**\u0026rsquo;\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eOviposition\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e312.919\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e104.306\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2.519\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.060\u0026lsquo;*\u0026rsquo;\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ePostoviposition\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e85.119\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e28.373\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e10.715\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.000***\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eOffspring/day\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e205.626\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e68.542\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.811\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.147\u0026lsquo;.\u0026rsquo;\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e "},{"header":"Discussion","content":"\u003cp\u003eIn the ecosystem, insects are not subjected to constant or alternating temperatures. However, laboratory conditions can provide valuable insight into the population dynamics of aphids. The findings reported here clearly show the effects of temperature on the development time, death rate, longevity, and fecundity of \u003cem\u003eA. craccivora\u003c/em\u003e. Aphid species are small ectothermic insects that are strongly related to temperature. Recall that temperature influences multiple aspects of insect biology, such as the metabolic system and developmental rate, as well as the timing and level of insect activities. Temperatures beyond the thresholds of development for a given insect can slow growth, and extreme temperatures can kill members of the insect population. It is logical that temperature fluctuations can cause changes in insect biology. In this study, we have discussed how temperature alternations affect insect biology. Several biological parameters of \u003cem\u003eAphis craccivora\u003c/em\u003e were also evaluated. The maximum temperature for the development of \u003cem\u003eA. craccivora\u003c/em\u003e was 28\u0026deg;C (mean\u0026thinsp;=\u0026thinsp;5.0 days; F\u0026thinsp;=\u0026thinsp;81.78; P\u0026thinsp;=\u0026thinsp;0.05), but other studies have reported a temperature of 30\u0026deg;C (Cho et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Kuo, M. H. \u0026amp; Chen, 2004) and a temperature of 29.4\u0026deg;C (Berberet et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Gir\u0026atilde;o et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) on bean plants. These results correlate with the degree to which ectothermic animals that develop under warm conditions tend to grow faster and mature earlier. On the other hand, these animals are slower in maturation than are similar animals that develop under cool conditions. The preoviposition period, during which the temperature is held constant and gradually decreases, delays the occurrence of fecundity. The temperature increases from 16\u0026deg;C to 24\u0026deg;C, similar to the 16\u0026deg;C and 25\u0026deg;C reported by (Cho et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Gir\u0026atilde;o et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The number of oviposition days strongly depends on the area in which the insect is evaluated by using food and weather resources. Normally, at both low- and high-tolerance temperatures, insects can take a long time to reproduce, for example, from 16\u0026deg;C to 28\u0026deg;C (mean\u0026thinsp;=\u0026thinsp;8.95, 5.07; F\u0026thinsp;=\u0026thinsp;2.519; P\u0026thinsp;=\u0026thinsp;0.05). The same tendency at 15, 20, and 25\u0026deg;C was reported by (Cho and et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) and at 18, 22, 25, and 28\u0026deg;C. (Gir\u0026atilde;o et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe postoviposition period was 16\u0026deg;C (mean day\u0026thinsp;=\u0026thinsp;2.05; F\u0026thinsp;=\u0026thinsp;2.519; P\u0026thinsp;=\u0026thinsp;0.05) if it was below the lowest relative temperature for good reproduction; during this time, the duration after the offspring period became longer than the adaptive temperatures reported at 18, 22, 25, and 28\u0026deg;C (Gir\u0026atilde;o, et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) (mean days\u0026thinsp;=\u0026thinsp;1.2, 0.8, 0.8, and 0.2, respectively). Longevity increases inversely with decreasing temperature, as we can see at 16\u0026deg;C (22.2 days) and 28\u0026deg;C (10.8 days); however, in the literature, approximately the same constant temperatures were reported at 18\u0026deg;C and 28\u0026deg;C by (Gir\u0026atilde;o et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eHowever, there were differences between the results presented in this research within the life span of \u003cem\u003eA. craccivora\u003c/em\u003e. The temperature regimes exhibited significant differences from the highest to lowest temperature at 28\u0026deg;C and 16\u0026deg;C (mean/day) (5.8 and 11.6, F\u0026thinsp;=\u0026thinsp;4.340; P\u0026thinsp;=\u0026thinsp;0.05, respectively). The mean values between the life span temperatures were similar: at 28\u0026deg;C, the lowest occurred at 16\u0026deg;C, and the highest occurred at 28\u0026deg;C. These findings revealed that there were no significant differences. These results indicate that these temperatures are amenable to survival for aphid species. (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). In general, these observations established the inverse proportional relationship between oviposition periods and immature stages produced per female per day in the temperature range of 16 to 28\u0026deg;C, in contrast to the results presented by(Berberet et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Gir\u0026atilde;o et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAccording to these results, there are many possible adaptations between several studied aphid populations. For example, the variations observed in thermal constant development based on the suitable temperature for black aphids on the bean might be explained in part by the functional theory of life, which relates to the highest value for a development threshold and the smallest value for total thermal requirement, as expected, for species more adapted to tropical regions and temperate regions (Trudgill \u0026amp; Perry, \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Brown et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1995\u003c/span\u003e).\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eConflict of interest\u0026nbsp;\u003c/strong\u003eThe authors declare that they have no conflicts of interest.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u0026nbsp;Ethical approval\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;This article does not contain any studies with human participants performed by any of the authors.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgments\u0026nbsp;\u003c/strong\u003eWe would like to thank the team management of the Project Development and Coordination Unit. This research was part of the Master\u0026rsquo;s degree thesis of Rochelyn DONA and was funded by the Project Development and Coordination Unit of Cukurova University (Grant Code: FYL-2020-12688).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u0026nbsp;\u003c/strong\u003eThis study was supported by Cukurova University. (Grants Code: FYL-2020-12688).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthors \u0026lsquo;contributions\u0026nbsp;\u003c/strong\u003eRD and SS conceived, designed and performed research. Both of the authors analyzed data, and wrote the manuscript. Both authors read and aprouved the manuscript.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAldryhim Y, Khalil A (1993) Influence of temperature and day length on population development of \u003cem\u003eAphis gossypii\u003c/em\u003e on \u003cem\u003eCucurbita pepo\u003c/em\u003e. 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J Appl Ecol, 431\u0026ndash;438\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCho JR, Kim JH, Choi BR, Seo BY, Kim KH, Ji CW, Ahn JJ (2018) Thermal effects on the development, fecundity and life table parameters of \u003cem\u003eAphis craccivora\u003c/em\u003e Koch (Hemiptera: Aphididae) on yardlong bean (Vigna unguiculata subsp. sesquipedalis (L). Korean J Appl Entomol 57:261\u0026ndash;269\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDedryver CA, Le Ralec A, Fabre F (2010) The conflicting relationships between aphids and men: a review of aphid damage and control strategies. CR Biol 333:539\u0026ndash;553\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eEbert TA, Cartwright B (1997) Biology and ecology of \u003cem\u003eAphis gossypii\u003c/em\u003e Glover (Homoptera: aphididae). 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Serie A: Seminaires Mediterraneens\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZhaozhi L, Likai F, Guizhen G, Ling-Ling G, Han P, Sharma S, Zalucki MP (2017) Differences in the high-temperature tolerance of \u003cem\u003eAphis craccivora\u003c/em\u003e (Hemiptera: Aphididae) on cotton and soybean: implications for ecological niche switching among hosts. Appl Entomol Zool 52:9\u0026ndash;18\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e\n"}],"fulltextSource":"","fullText":"","funders":[{"identity":"9f70c415-1ed1-4311-b537-230fd89b0719","identifier":"10.13039/501100002964","name":"Çukurova Üniversitesi","awardNumber":"Grants Code: FYL-2020-12688","order_by":0}],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":true,"highlight":"","institution":"Cukurova University","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Adult longevity, life table, nymphal stage, nonlinear function, thermal resistance","lastPublishedDoi":"10.21203/rs.3.rs-4960855/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4960855/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe cowpea aphid \u003cem\u003eAphis craccivora\u003c/em\u003e is a polyphagous species that has spread worldwide. The aim of this study was to investigate the thermal effects on the developmental period, longevity, and fecundity of apterous females of \u003cem\u003eA. craccivora\u003c/em\u003e. The experiment was conducted under the effects of four temperature regimes, which included 16\u0026deg;C, 20\u0026deg;C, 24\u0026deg;C, and 28\u0026deg;C, with 65\u0026thinsp;\u0026plusmn;\u0026thinsp;5% relative humidity (RH) and a photoperiod of long-day 16:8 (L:D) h. Afterwards, the nymphs developed successfully until the adult stage at all temperature regimens. The developmental period of immature \u003cem\u003eA. craccivora\u003c/em\u003e ranged from 10.6 days at 16\u0026deg;C to 5.0 days at 28\u0026deg;C. The nymph viability and survival were greater at 24\u0026deg;C than at the other temperatures. However, at a constant temperature of 28\u0026deg;C, the mortality rate was greater than that of the other species at the immature stage of \u003cem\u003eA. craccivora\u003c/em\u003e. The lower developmental threshold for cowpea aphids was estimated at 1.77\u0026deg;C and 66.79 degree-days (DD) at the first instar until adulthood. The average longevity of adult females decreased from 22.2 days at 16\u0026deg;C to 10.8 days at 28\u0026deg;C. The net reproduction rate per female was 46.97 at 24\u0026deg;C and 26.93 to 28\u0026deg;C. The largest intrinsic rates of increase (\u003cem\u003er\u003c/em\u003e\u003csub\u003e\u003cem\u003em\u003c/em\u003e\u003c/sub\u003e= 0.367) occurred at 28\u0026deg;C, and the smallest occurred at 16\u0026deg;C (\u003cem\u003er\u003c/em\u003e\u003csub\u003e\u003cem\u003em\u003c/em\u003e\u003c/sub\u003e= 0.177). It was obvious that temperatures greater than 28\u0026deg;C resulted in good development, increased mortality at the nymphal stage, reduced adult longevity, and diminished fecundity. The optimal growth temperature for \u003cem\u003eA. craccivora\u003c/em\u003e was 20\u0026deg;C-24\u0026deg;C.\u003c/p\u003e","manuscriptTitle":"Thermal effects on the biological parameters of bean Aphis craccivora (Hemiptera: Aphididae)","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-08-26 07:53:08","doi":"10.21203/rs.3.rs-4960855/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"b7f711c4-255b-4626-bedb-aff39ae78255","owner":[],"postedDate":"August 26th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":36449638,"name":"Entomology"},{"id":36449639,"name":"Behavioral Ecology"},{"id":36449640,"name":"Developmental Biology"},{"id":36449641,"name":"Population Biology"}],"tags":[],"updatedAt":"2024-08-26T07:53:08+00:00","versionOfRecord":[],"versionCreatedAt":"2024-08-26 07:53:08","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-4960855","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4960855","identity":"rs-4960855","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}