Thermal effects on the growth and developmental time of bean Aphis fabae (Hemiptera: Aphididae)

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Abstract The aim of this study was to evaluate the thermal effects on the biological parameters of Aphis fabae Scopoli (Hemiptera: Aphididae). The developmental times, survival, and reproductive data were collected for Aphis fabae reared on detached bean leaves (Phaseolus vulgaris L.) ‘pinto beans at five temperatures regimes (12°C, 16°C, 20°C, 24°C and 28°C), 65% relative humidity (RH), and a photoperiod of 16:8 (L: D) h. The developmental times of immature stages ranged from 16,65 days at 12°C to 5.70 days at 24°C, but a slight increase again at 28°C (6.62 days). This study presented the optimal developmental threshold for A. fabae slightly to 24°C. The average longevity of mature females significantly decreased from 42.32 days at 12°C to 16.12 days at 28°C. The reproduction rate per female was 62.27 individuals at 16°C and 12.72 individuals at 28°C. The mean generation period of the population ranged from 29.24 at 12°C to 11.50 at 28°C. The highest intrinsic rate of increase (r = 0.41) was recorded at 24°C, the lowest at 12°C (r = 0.15). It was evident that temperatures over 28°C augmented the development time, accelerated the death ratio of the nymphal stages, Shrunk Adult longevity, and reduced fecundity. The optimal range of temperature for population growth of A. fabae on bean was 16°C-24°C according to this study.
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Thermal effects on the growth and developmental time of bean Aphis fabae (Hemiptera: Aphididae) | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Thermal effects on the growth and developmental time of bean Aphis fabae (Hemiptera: Aphididae) Rochelyn DONA, Serdar Satar This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-5220155/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 5 You are reading this latest preprint version Abstract The aim of this study was to evaluate the thermal effects on the biological parameters of Aphis fabae Scopoli (Hemiptera: Aphididae). The developmental times, survival, and reproductive data were collected for Aphis fabae reared on detached bean leaves ( Phaseolus vulgaris L. ) ‘pinto beans at five temperatures regimes (12°C, 16°C, 20°C, 24°C and 28°C), 65% relative humidity (RH), and a photoperiod of 16:8 (L: D) h. The developmental times of immature stages ranged from 16,65 days at 12°C to 5.70 days at 24°C, but a slight increase again at 28°C (6.62 days). This study presented the optimal developmental threshold for A. fabae slightly to 24°C. The average longevity of mature females significantly decreased from 42.32 days at 12°C to 16.12 days at 28°C. The reproduction rate per female was 62.27 individuals at 16°C and 12.72 individuals at 28°C. The mean generation period of the population ranged from 29.24 at 12°C to 11.50 at 28°C. The highest intrinsic rate of increase ( r = 0.41) was recorded at 24°C, the lowest at 12°C ( r = 0.15). It was evident that temperatures over 28°C augmented the development time, accelerated the death ratio of the nymphal stages, Shrunk Adult longevity, and reduced fecundity. The optimal range of temperature for population growth of A. fabae on bean was 16°C-24°C according to this study. Developmental time intrinsic rate optimal range reproduction period temperature regimes Figures Figure 1 Figure 2 Figure 3 Introduction Aphids are considered a serious pest in agriculture crops and are characterized by their rapid development time with a huge potential of reproduction. They do not only cause direct damages to their host plants, but they also contribute as vector transmission viruses of plant pathogens. Their taxonomy was discussed by (Blackman and Eastop, 2007). In this study, the authors reported that the taxa of the fabae group are morphologically difficult to separate (Tosh et al., 2004). A. fabae , because it is typically anholocyclic (Barbagallo and Stroyan, 1982). The heteroecious and monoecious holocycles that occur on mountains are also known (Blackman and Eastop, 2006). The black aphid is considered as the most important pest attacking faba bean ( Vicia faba L.), around the world it is cultivated (Adabi and Zamani, 2010; Singh and Sharma, 2016; Blackman and Eastop, 2016 ; Webster, 2012 ; Béji, and Makni 2015; Al-Antary and Thaliji, 2007; Mahmoudi et al., 2010 ). In the eastern Mediterranean region is known as one of the most nuisible in the Yalova Province of Turkey (Kuloğlu and Özder, 2017 ; Akca and Chi, 2015). This insect could be found in many field crops, such as beans, tomatoes, potatoes, tobacco, and wild ornamental plants. For example, from Fabaceae included over the range of 200 leguminous plants and infested all their parts all over the world (Völkl and Stechmann 1998 ; Barnea et al., 2005; Izzet et al., 2015). From the genus of beans, Mullussurmuletus L. (fabales: Fabaceae), phenological stages; such as growth and development can be disturbed. A. fabae has been considered as a pest numerous on bean fields at this time presents to create serious problems in our ecosystem in reducing the growth and development of crop yield. Host plants are damaged either directly by aphid feeding or indirectly by the transmission of viruses and excretion of honeydew (MILLS, 1989; SCHEPERS, 1989). Previous studies assessed the flowers can abort and the honeydew secreted by the Aphid on the leaves provoke a reduction of photosynthetic process in causing sooty mold in response to A. fabae saliva and extraction of liquid sugar-rich sticky from the plant phloem (Nuessly et al. 2004, Izzet et al. 2015). The insect physiologies and their fitness are significantly influenced by the temperature fluctuation they encounter (Tiring, et al 2023 ). Insects could therefore be sensitive within short periods to climate change at the population level, such that the fingerprints of even early stages of climate change could be seen on insect population density and the number of generations. Insects are ectothermic organisms, for this reason, the temperature regimes can provoke a major impact on their development, growth, survival, reproduction, and behavior (Harvey., et al 2020). Consequently, insect pests such as aphids and all other arthropods depending on the weather conditions may favorite rapidly their decreasing, increasing development, and even death. Some studies reported the development rate of the insects at different temperatures focused only on the level temperature-developmental rate of A. fabae (Frazer, 1972 ; Mahmoudi and Sendi, 2010; Sabaghi and Hosseini, 2011; Dixon and Wratten, 1971 ; O'Doherty, 1986 ;). These articles are related to a simple variable of insect development and growth. But this one has been conducted on demography and population projection of Aphis fabae with additional comments on life table research criteria (Akca, Ayvaz, Yazici, Smith, and Chi ( 2015 ). The present study was designed to provide data on A. fabae at five constant temperatures and analyzed them using growth and development time. Finally, according to the global warming, understanding biological responses of black bean aphid on the effect of alternating temperatures that may use for IPM decision making on the level of development of bean pest. Materials and methods Plant culture. Pinto bean, Phaseolus vulgaris L . was cultured into the plant home 125 plants were placed with an alternation of one week each other in the goal having on time the fresh leaves for feeding the insects. The condition of the home plant was consistenly set up at 25 ± 1°C and 65 ± 10% RH and a photoperiod of 16h artificial light around 10000 Lux. (Fig. 1 ). Insect culture. The black bean aphid, A. fabae was collected on the bean plant, Vicia fabae (Sevilla) properly from the experimental field of plant protection, department of agriculture at Çukurova University, Balcali/Adana, and Turkey. More than > 50 aphids at the adult stage were reared inside of 5 cages on pinto beans ( P. vulgaris L .) in laboratory conditions at 24 ± 1°C, 65 ± 5% RH, and a photoperiod of 16:8 (L: D) h about 5 generations before individuals were used in the experiments. (Fig. 1 ). Experimental procedures This study was randomly conducted by selected the females apterous from the stock cage culture and individually transferred to the undersurface of bean leaves in the plastic Petri dishes (both 5_cm of diameter) at each level of temperature. From them, 10 (ten) master Petri dishes as stock have been placed inside the incubators. After 3 generations, 40 Petri dishes were prepared with a wetted cotton pad (0.5 cm) and placed under the leaves covered all the surface of wetted cotton to avoid them from drying. After that, 40 nymph stages were taken with a paintbrush carefully from the master stock to the new others. The cotton wool in the Petri dishes was wetted daily and every 3–5 days the aphids were transferred to new cotton leaf discs. The fresh used leaves were taken from the plants at the home condition and brought to the laboratory. The experiments were conducted on five constant temperatures of 12, 16, 20, 24 and 28 ± 1°C, 60 ± 5% RH and a photoperiod of 16:8 (L: D) 24h. Each temperature respectively was started with 40 newborn nymphs. Every 24 h the nymphal development was recorded until the adult stage. After the adult period, the number of newly born and survival produced by the mother were registered until the death of all adults. Statistical Analysis. Data concerning developmental time, fecundity, longevity, survivorship, intrinsic rate of natural increase, and the jackknife estimate of rm for the cotton cultivar experiments were evaluated using the ANOVA. Fecundity and longevity data were log10 transformed to stabilize the variance before analysis. With the ANOVA application, using the statistical software SPSS 11.5 (SPSS Inc., Chicago, IL) (Green et al. 2000), differences between the means were evaluated using the least significant difference (LSD) test at P _ 0.05. Differences classification of variables, developmental time and reproductive performance were subjected to descriptive statistical analysis (one way-ANOVA) for example, the different phrenological stages of A. fabae , including development times for nymphal molts, reproduction periods and determining lifetime of adults on each constant temperature. Furthermore, the mention of the calculation of life table’s parameters at five temperature regimes is completely randomized design (CRD). Eventually, the multiple comparison data of averages were done in testing the significant difference whe n they have existed using Tukey’s HSD multiple range test (P = 05) the calculations were realized by using Spss. For each constant temperatures curves were calculated employing a product limit technique. Population growth rates were computed from the equation of Lotka (23): 1 = Σ e-r*x lx * mx (1) In which: x = age in days, r = intrinsic rate of increase, Lx = age-specific survival, mx = age-specific number of female offspring. After "r" was computed for the original data (rall), differences among rm-values were tested for significance by estimating variances through the jackknife method (24). The jackknife pseudo-value "rj" was calculated for the "n" samples using the following equation: rj = n * rall - (n-1) *ri (2) The mean of "n" jackknife pseudo-values for each treatment was subjected to analysis of variance. The scheffé test was used to compare mean growth rates for different temperature regimes (p = 0.05). Because low probability levels were used, there was no concern about inflation of experiment-wise error rates (25). Each of the abovementioned analyses were conducted using the Statgraphics software package. Results The developmental time of A. fabae at five constant temperatures is summarized below (Table 1 ). The results showed that the different instar stages significantly decreased with increasing constant temperatures. The first-instar stage ranged from 1.27 days at 20°C to 3 days at 12°C and there is no significant difference with 1.55 days at 24°C, 1.70 days at 28°C (F = 84.03; P = 0.05). For the second instar stage there is significant difference between low and high constant temperature ranged from 1.37 days at 28°C to 3.30 days to 12°C (F = 63.68; P = 0.05). At the third-period from 1.30 days 24°C to 5.80 days at 12°C, but a slight difference at 28°C with 1.75 days (F = 96.65; P = 0.05). And at fourth instar stage from 1.52 days at 24°C to 4.55 days to 12°C (F = 52.73; P = 0.05). Moreover, for the total development time (TDT), the same pattern was presented. The instar stage was shorter at high temperature and longer at low temperature. Significant increase was found for TDT at 12°C and 16°C (16.65 and 11.58 days respectively) compared to 20°C, 24°C and 28°C (6.22, 5.70, and 6.62 days respectively) that are statistically similar (Table.1). Concerning the variation days at 28°C treatment. High mortality was observed during immature stages causing a slight difference between the results 24°C and 28°C. The pre-reproduction period showed a significance variation between temperature regimes, increasing at high temperature and increased at low temperature (0.27 days at 28°C to 1.10 days to 16°C F = 13.54; P = 0.05). Table 1 Development times (days Mean ± SEM) of Aphis fabae on bean ( Phaseolus vulgaris L.), at five constant temperatures, 65 ± 5% RH, and a photoperiod of 16:00(L: D) h. Temp(°c) N1 (day) N2 (day) N3 (day) N4 (day) Total development (day) Pre-reproduction (day) 12°C 3.00 ± 0.0a 3.30 ± 0.1a 5.80 ± 0.3a 4.55 ± 0.2a 16.65 ± 0.2a 0.97 ± 0.1ab 16°C 2.40 ± 0.1b 3.15 ± 0.1a 2.45 ± 0.1b 3.70 ± 0.2b 11.58 ± 0.4b 1.10 ± 0.0a 20°C 1.27 ± 0.0d 1.57 ± 0.0b 1.80 ± 0.1bc 1.57 ± 0.1c 6.22 ± 0.1c 0.32 ± 0.0cd 24°C 1.55 ± 0.0cd 1.41 ± 0.0b 1.30 ± 0.0c 1.52 ± 0.0c 5.70 ± 0.1c 0.67 ± 0.1bc 28°C 1.70 ± 0.0c 1.37 ± 0.0b 1.75 ± 0.0bc 1.80 ± 0.1c 6.62 ± 0.1c 0.27 ± 0.1d Significant differences between means ( * P < 0.05 and ** P < 0.01) are expressed by different letters (a–d). The letters compare values in the same column. A linear regression analysis was applied to the developmental level of temperature from 12°C to 28°C range. Within the selected temperature range the developmental rates (r (t) ) of A. craccivora increased linearly with the increasing of the temperature (Figure. 2). The theoretical developmental threshold of A craccivora was recorded as 1.75°C and required 66.66 degree-days for completing all the immature stages to become adult. The number of days of reproduction was significantly longer at low temperature, at 12°C it was 23.95 days (F = 31.54: P = 0.05) and decreased statistically at higher temperature (28°C; 6.67 days) (Table 2 ). The reproduction period was statistically the same for all temperature regimes and ranged from 0.92 days at 16°C to 2.05 days at 20°C (F = 2.09; P = 0.05). It was shorter at lower temperature (Table 2 ). Longevity was significantly longer at low temperature (42.33 days at 12°C and 32.6 days at 16°C compared to the others. However, 20°C (22.67 days) was statistically similar to 24°C (20.05 days) and the latter statistically similar to 28°C (16.12 days) (Table 2 ). The life span of A. fabae days was significantly longer at low temperature 12°C 25.67days and decreased statistically at higher temperature (28°C 9.50 days F = 21.50; P = 0.05), (Table 2 ). Table 2 Reproduction, post- reproduction, Longevity, life span and number of offspring of Aphis fabae adult female individuals (mean ± SE). Temp (°C) Reproduction (day) Post- reproduction (day) Longevity (day) Life span (day) Number of offspring (Aphid) 12°C 23.95 ± 0.8a 0.95 ± 0.21 42.32 ± 0.8a 25.67 ± 0.9 a 49.57 ± 2.6a 16°C 18.87 ± 1.5b 0.92 ± 0.24 32.60 ± 1.6b 21.02 ± 1.5ab 62.27 ± 5.2a 20°C 14.00 ± 1.3c 2.05 ± 0.45 22.67 ± 1.6c 16.45 ± 1.6bc 55.00 ± 4.6a 24°C 11.47 ± 1.1c 1.42 ± 0.47 20.05 ± 1.3cd 14.35 ± 1.3cd 48.67 ± 4.5a 28°C 6.67 ± 0.8d 2.00 ± 0.40 16.12 ± 1.2d 9.50 ± 1.2d 16.72 ± 2.5b Significant differences between means ( * P < 0.05 and ** P < 0.01) are expressed by different letters (a–d). The letters compare values in the same column. Virtually at the high level of temperatures, the mother aphids were less survived. The number of newborns per days, their size was reduced and early died at 28°C (Fig. 3 ). This study was resulted that with an increasing of constant temperatures, death ratio of the immature populations was more effectively. For example, at 28°C, there is 25% of death compared to the lowest temperature regimes The number of generation are produced per (To) A. fabae at 12°C were 29.14 generations while at 28°C 11.50 generations (Table 3 ). The net reproduction (Ro) was higher at 16°C (62.27 aphids/aphid) while at low at 28°C (16.72 aphids/aphid) (Table 3 ). The offspring per reproduction days was significantly longer at 24°C 4.84 aphids/days compared to others. A. fabae at middle temperatures was well developed and showed the highest population growth per capita rate across the intrinsic rate of increase ranged at 20°C and 24°C (0.35 and 0.36 aphids/aphid per days) and the lowest at 12°C (0.15 aphids/aphid per day). The number of offspring was significantly different at 28°C 16. 72 aphids compared to the others. Table 3 Generation time (T0), net reproduction (Ro), and intrinsic rate of increase (rm) of Aphis fabae on bean leaf discs at five temperatures levels Temp (°C) Sample size(N) Female adults(N) Death ratio (%) Generation time(T 0 ) Reproduction rate (R 0 ) Offspring per production day (Mean ± SEM) Intrinsic rate increase(aphis/aphid/d) 12°C 40 40 0 29.14 50.96 2.08 ± 0.7b 0.15 16°C 40 36 10 23.17 62.27 2.08 ± 0.1b 0.21 20°C 40 38 5 14.40 55 3.99 ± 0.2 ab 0.35 24°C 40 38 5 12.97 48.68 4.84 ± 0.5a 0.36 28°C 40 30 25 11.50 16.72 2.08 ± 0.3b 0.27 Significant differences between means ( * P < 0.05 and ** P < 0.01) are expressed by different letters (a–d). The letters compare values in the same column. Discussion Temperature affects the development as well as growth time; consequently, the evolution and dynamics of insect populations in the field are dictated by ambient temperature. For this reason, there is a considerable interest for a long time in the temperature relationships of development time, reproduction, nymphal mortality rate, longevity, and fecundity of A. fabae. According to the outcomes of this study transparently, showed that at five constant temperatures on biological development of A. faba e (Table 1 , 2 , 3 ). The different nymphal stages and total development significantly decreased from 12°C until 24°C with increasing constant temperature, however from 24°C to 28°C of the total developmental was augmented again. According to some similar studies were reported by (Tofangsazi et al., 2012 ; Akca et al., 2015 ). Insect cultured at the temperatures over the threshold develop more slowly than others that reared below favorable conditions (Satar et al., 1999 and 2005 ; Kersting et al., 1999 ; Dona and Satar ( 2024 ). Another observation has been related by Olmez et al., 2003; Maelzer, 1977) who reported that the developmental rate of M. rosae was retarded at temperatures above 25°C to 32°C. This study presented the same reality for A. fabae on faba bean at 28°C showing a significant increase in developmental time evaluated with that at 24°C. Therefore, only at 20°C in the second-instar stage, a slight increase has been presented comparing with 24°C. The percentage of mortality at 28°C has been shown highest elevation death, while at 12°C, the smallest it resists at hundred percent (100%). According to the physiology of A. fabae , its adaptation reputes more for cold temperatures than high. Although it occurs in tropical areas of Africa, it is not a very serious pest in warm environments (Farhadi et al., 2011 ). Generally, aphids at lower temperatures need to take more developmental time before they start to produce nymphs (Table 1 ). reproduction period, the duration of reproduction of A. fabae clearly showed as much as decreasing over threshold temperature limit take more time for reproducing. For example, at 12°C 23.95 days to 28°C 6.67days. The post- reproduction period didn’t show any significant differences between the temperatures (Table.2) Longevity and life span from some previous studies reported that decreased constant temperature engenders an increasing period. It’s a common phenomenon from various aphid species (Morgan et al., 2001 ; Wang & Tsai, 2000; Collins & Leather, 2001; Satar & Yokomi, 2002). Another observation has been related by (Olmez et al., 2003; Akca et al., 2015 and Maelzer. 1977) who reported that the developmental rate of M. rosae was retarded at temperatures above 25°C. While in our study over 24°C similar results were presented. The intrinsic rate of natural increase ( r ) is a key an indicator that can be demonstrated as the most favorable temperature level for better population growth because it comes from all over impacts caused by the temperature on the life table of a population. The population reared from 12 to 24°C, this last presented the highest ( r- value compare with others ( r = 0.36 aphids/aphid per day) because it the most rapid development, the highest survival of nymphal stages in an average of number 4.80 nymphs of daily progeny and proved as long as r -value exposed at an alternate temperature couldn’t be subjected as the highest net production rate per female for A. fabae , but in this case with an increasing of constant temperature 24°C to 16°C ( Ro = 48.68 to 62.27). A similar result has been reported by (Akca et al., 2015 ) gradually decreasing from 15 to 25°C ( r = 0.43 d-1) and the opposite for ( Ro = 90.44 to 77.46) increased. According to those results from the smallest to the biggest the most suitable temperature of A, fabae ranged 16 to 24°C, otherwise, its biology can be affected greatly. On the other hand, the host variety also can play a major role in aphid biology. Related by (Fernandez-Quintanilla et al., 2002) about a range of life parameters of A. fabae on weed species in winter and summer as 0.14 to 0.36 days (the mean unit used in this article was nymphs per female per days). Declarations Conflict of Interest The authors declare that they have no conflict 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 Msc of Rochelyn DONA was funded by Project Development and Coordination Unit from Çukurova University (Grants Code: FYL-2020-12688). 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Journal of Asia-Pacific Entomology , 13 (2), 111-116. Mahmoudi, M., Sahragard, A., & Sendi, J. J. (2010). Foraging efficiency of Lysiphlebus fabarum Marshall (Hymenoptera: Aphidiidae) parasitizing the black bean aphid, Aphis fabae Scopoli (Hemiptera: Aphididae), under laboratory conditions. Journal of Asia-Pacific Entomology , 13 (2), 111-116. Morgan, D., Walters, K. F. A., & Aegerter, J. N. (2001). Effect of temperature and cultivar on pea aphid, Acyrthosiphon pisum (Hemiptera: Aphididae) life history. Bulletin of entomological research , 91 (1), 47. O'Doherty, R. (1986). Cold hardiness of laboratory-maintained and seasonally-collected populations of the black bean aphid, Aphis fabae Scopoli (Hemiptera: Aphididae). Bulletin of entomological research , 76 (3), 367-374. Sabaghi, R., Sahragard, A., & Hosseini, R. (2011). Functional and numerical responses of Scymnus syriacus Marseul (Coleoptera: Coccinellidae) to the black bean aphid, Aphis fabae Scopoli (Hemiptera: Aphididae) under laboratory conditions. Journal of Plant Protection Research , 51 (4), 423-428. Satar, S., Kersting, U., & Uygun, N. (1999). Development and fecundity of Aphis gossypii Glover (Homoptera: Aphididae) on three Malvaceae hosts. Turkish Journal of Agriculture and Forestry , 23 (6), 637-644. Satar, S., Kersting, U., & Uygun, N. (2005). Effect of temperature on development and fecundity of Aphis gossypii Glover (Homoptera: Aphididae) on cucumber. Journal of Pest Science , 78 (3), 133-137. Singh, R., Singh, G., Singh, K., & Sharma, A. (2016). Biodiversity of Aphids (Insecta: Homoptera: Aphididae) Infesting Legumes (Angiospermae: Fabales: Fabaceae) in India. International Journal of Research Studies in Zoology (IJRSZ) , 2 (1), 30-44. Tiring, G., Ada, M., Ada, M., Dona, R., & Satar, S. (2023). The effect of climate on the population of Grapholita molesta (Busck) (Lepidoptera: Tortricidae) in Mersin, Türkiye. Erwerbs-obstbau , 65 (6), 2289-2297. Tofangsazi, N., Kheradmand, K., Shahrokhi, S., & Talebi, A. (2012). Effect of different constant temperatures on biology of Schizaphis graminum (rondani) (Hemiptera: aphididae) on barley, Hordeum vulgare L. (poaceae) in Iran. Journal of plant protection research . Völkl, W., & Stechmann, D. H. (1998). Parasitism of the black bean aphid (Aphis fabae) by Lysiphlebus fabarum (Hym., Aphidiidae): the influence of host plant and habitat 1. Journal of Applied Entomology , 122 (1‐5), 201-206. Webster, B. E. N. (2012). The role of olfaction in aphid host location. Physiological Entomology , 37 (1), 10-18. Béji, B., Bouhachem-Boukhris, S., Bouktila, D., Mezghani-Khémakhem, M., Rezgui, S., Kharrat, M., & Makni, H. (2015). Identification of sources of resistance to the black bean aphid, Aphis fabae Scopoli, in Faba bean, Vicia faba L., accessions. Journal of Crop Protection , 4 (2), 217-224. Cite Share Download PDF Status: Under Review Version 1 posted Editorial decision: Major revisions 03 Jun, 2025 Reviewers agreed at journal 14 May, 2025 Reviewers invited by journal 09 May, 2025 Editor assigned by journal 17 Oct, 2024 First submitted to journal 15 Oct, 2024 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. We do this by developing innovative software and high quality services for the global research community. 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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-5220155","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":454313798,"identity":"79842fb5-9bc5-41ab-bd36-94d412028687","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 Universitesi Ziraat Fakultesi","correspondingAuthor":true,"prefix":"","firstName":"Rochelyn","middleName":"","lastName":"DONA","suffix":""},{"id":454313799,"identity":"1beb1828-38e8-4602-98bc-9f7b027ce979","order_by":1,"name":"Serdar Satar","email":"","orcid":"","institution":"Cukurova Universitesi Ziraat Fakultesi","correspondingAuthor":false,"prefix":"","firstName":"Serdar","middleName":"","lastName":"Satar","suffix":""}],"badges":[],"createdAt":"2024-10-07 18:34:29","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-5220155/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-5220155/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":82740876,"identity":"c49ca9c9-9ff0-46fe-9942-d36a0cc40929","added_by":"auto","created_at":"2025-05-14 17:03:07","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":130806,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eA\u003c/strong\u003e) Home condition of plantation, \u003cstrong\u003eb)\u003c/strong\u003e incubator biological inside it, insect cultured and 24°C \u003cstrong\u003ec) \u003c/strong\u003enymphal counted\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-5220155/v1/2a695a847250bbf645df36fa.png"},{"id":82740878,"identity":"c206b3ca-72d8-4033-9480-e78a393990dc","added_by":"auto","created_at":"2025-05-14 17:03:07","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":41901,"visible":true,"origin":"","legend":"\u003cp\u003eDevelopmental rate (r) of \u003cem\u003eA. fabae\u003c/em\u003e at five constants (blue squares) and all of them alternated (variation cycle), analysis of the linear regression line of the developmental rate ranged from12°C to 28°C. The means alternating temperatures were used to calculate the regression analysis.\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-5220155/v1/d898be4c4bbf29a7b7510a2d.png"},{"id":82741718,"identity":"0d2110f5-1811-4bb0-a003-885e56a8bb10","added_by":"auto","created_at":"2025-05-14 17:11:07","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":193163,"visible":true,"origin":"","legend":"\u003cp\u003eAge- specific survival rate (lx) and fecundity (mx) of Aphis fabae reared on faba bean leaves \u003cem\u003ePhaseolus vulgaris \u003c/em\u003eL. at five constant temperatures\u003c/p\u003e","description":"","filename":"floatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-5220155/v1/446f6147b55245a080d8551c.png"},{"id":82742048,"identity":"ff0d095b-2287-4e9c-ba1d-fb79b6d24765","added_by":"auto","created_at":"2025-05-14 17:19:08","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1015576,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-5220155/v1/e30e5874-dfc4-4a88-8134-c46997024026.pdf"}],"financialInterests":"","formattedTitle":"\u003cp\u003eThermal effects on the growth and developmental time of bean \u003cem\u003eAphis\u003c/em\u003e fabae (Hemiptera: Aphididae)\u003c/p\u003e","fulltext":[{"header":"Introduction","content":"\u003cp\u003eAphids are considered a serious pest in agriculture crops and are characterized by their rapid development time with a huge potential of reproduction. They do not only cause direct damages to their host plants, but they also contribute as vector transmission viruses of plant pathogens. Their taxonomy was discussed by (Blackman and Eastop, 2007). In this study, the authors reported that the taxa of the fabae group are morphologically difficult to separate (Tosh et al., 2004). \u003cem\u003eA. fabae\u003c/em\u003e, because it is typically anholocyclic (Barbagallo and Stroyan, 1982). The heteroecious and monoecious holocycles that occur on mountains are also known (Blackman and Eastop, 2006). The black aphid is considered as the most important pest attacking faba bean (\u003cem\u003eVicia faba\u003c/em\u003e L.), around the world it is cultivated (Adabi and Zamani, 2010; Singh and Sharma, 2016; Blackman and Eastop, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Webster, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; B\u0026eacute;ji, and Makni 2015; Al-Antary and Thaliji, 2007; Mahmoudi et al., \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). In the eastern Mediterranean region is known as one of the most nuisible in the Yalova Province of Turkey (Kuloğlu and \u0026Ouml;zder, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Akca and Chi, 2015).\u003c/p\u003e \u003cp\u003eThis insect could be found in many field crops, such as beans, tomatoes, potatoes, tobacco, and wild ornamental plants. For example, from Fabaceae included over the range of 200 leguminous plants and infested all their parts all over the world (V\u0026ouml;lkl and Stechmann \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Barnea et al., 2005; Izzet et al., 2015). From the genus of beans, \u003cem\u003eMullussurmuletus\u003c/em\u003e L. (fabales: Fabaceae), phenological stages; such as growth and development can be disturbed. \u003cem\u003eA. fabae\u003c/em\u003e has been considered as a pest numerous on bean fields at this time presents to create serious problems in our ecosystem in reducing the growth and development of crop yield. Host plants are damaged either directly by aphid feeding or indirectly by the transmission of viruses and excretion of honeydew (MILLS, 1989; SCHEPERS, 1989). Previous studies assessed the flowers can abort and the honeydew secreted by the Aphid on the leaves provoke a reduction of photosynthetic process in causing sooty mold in response to \u003cem\u003eA. fabae\u003c/em\u003e saliva and extraction of liquid sugar-rich sticky from the plant phloem (Nuessly et al. 2004, Izzet et al. 2015).\u003c/p\u003e \u003cp\u003eThe insect physiologies and their fitness are significantly influenced by the temperature fluctuation they encounter (Tiring, et al \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Insects could therefore be sensitive within short periods to climate change at the population level, such that the fingerprints of even early stages of climate change could be seen on insect population density and the number of generations. Insects are ectothermic organisms, for this reason, the temperature regimes can provoke a major impact on their development, growth, survival, reproduction, and behavior (Harvey., et al 2020). Consequently, insect pests such as aphids and all other arthropods depending on the weather conditions may favorite rapidly their decreasing, increasing development, and even death. Some studies reported the development rate of the insects at different temperatures focused only on the level temperature-developmental rate of \u003cem\u003eA. fabae\u003c/em\u003e (Frazer, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1972\u003c/span\u003e; Mahmoudi and Sendi, 2010; Sabaghi and Hosseini, 2011; Dixon and Wratten, \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1971\u003c/span\u003e; O'Doherty, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e1986\u003c/span\u003e ;). These articles are related to a simple variable of insect development and growth. But this one has been conducted on demography and population projection of Aphis fabae with additional comments on life table research criteria (Akca, Ayvaz, Yazici, Smith, and Chi (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). The present study was designed to provide data on \u003cem\u003eA. fabae\u003c/em\u003e at five constant temperatures and analyzed them using growth and development time. Finally, according to the global warming, understanding biological responses of black bean aphid on the effect of alternating temperatures that may use for IPM decision making on the level of development of bean pest.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cp\u003e \u003cb\u003ePlant culture.\u003c/b\u003e Pinto bean, \u003cem\u003ePhaseolus vulgaris L\u003c/em\u003e. was cultured into the plant home 125 plants were placed with an alternation of one week each other in the goal having on time the fresh leaves for feeding the insects. The condition of the home plant was consistenly set up at 25\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u0026deg;C and 65\u0026thinsp;\u0026plusmn;\u0026thinsp;10% RH and a photoperiod of 16h artificial light around 10000 Lux. (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eInsect culture.\u003c/b\u003e The black bean aphid, \u003cem\u003eA. fabae\u003c/em\u003e was collected on the bean plant, \u003cem\u003eVicia fabae\u003c/em\u003e (Sevilla) properly from the experimental field of plant protection, department of agriculture at \u0026Ccedil;ukurova University, Balcali/Adana, and Turkey. More than \u0026gt;\u0026thinsp;50 aphids at the adult stage were reared inside of 5 cages on pinto beans (\u003cem\u003eP. vulgaris L\u003c/em\u003e.) in laboratory conditions at 24\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u0026deg;C, 65\u0026thinsp;\u0026plusmn;\u0026thinsp;5% RH, and a photoperiod of 16:8 (L: D) h about 5 generations before individuals were used in the experiments. (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eExperimental procedures\u003c/h2\u003e \u003cp\u003eThis study was randomly conducted by selected the females apterous from the stock cage culture and individually transferred to the undersurface of bean leaves in the plastic Petri dishes (both 5_cm of diameter) at each level of temperature. From them, 10 (ten) master Petri dishes as stock have been placed inside the incubators. After 3 generations, 40 Petri dishes were prepared with a wetted cotton pad (0.5 cm) and placed under the leaves covered all the surface of wetted cotton to avoid them from drying. After that, 40 nymph stages were taken with a paintbrush carefully from the master stock to the new others. The cotton wool in the Petri dishes was wetted daily and every 3\u0026ndash;5 days the aphids were transferred to new cotton leaf discs. The fresh used leaves were taken from the plants at the home condition and brought to the laboratory.\u003c/p\u003e \u003cp\u003eThe experiments were conducted on five constant temperatures of 12, 16, 20, 24 and 28\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u0026deg;C, 60\u0026thinsp;\u0026plusmn;\u0026thinsp;5% RH and a photoperiod of 16:8 (L: D) 24h. Each temperature respectively was started with 40 newborn nymphs. Every 24 h the nymphal development was recorded until the adult stage. After the adult period, the number of newly born and survival produced by the mother were registered until the death of all adults.\u003c/p\u003e \u003cp\u003e \u003cb\u003eStatistical Analysis.\u003c/b\u003e Data concerning developmental time, fecundity, longevity, survivorship, intrinsic rate of natural increase, and the jackknife estimate of rm for the cotton cultivar experiments were evaluated using the ANOVA. Fecundity and longevity data were log10 transformed to stabilize the variance before analysis. With the ANOVA application, using the statistical software SPSS 11.5 (SPSS Inc., Chicago, IL) (Green et al. 2000), differences between the means were evaluated using the least significant difference (LSD) test at \u003cem\u003eP\u003c/em\u003e _ 0.05.\u003c/p\u003e \u003cp\u003eDifferences classification of variables, developmental time and reproductive performance were subjected to descriptive statistical analysis (one way-ANOVA) for example, the different phrenological stages of A. \u003cem\u003efabae\u003c/em\u003e, including development times for nymphal molts, reproduction periods and determining lifetime of adults on each constant temperature. Furthermore, the mention of the calculation of life table\u0026rsquo;s parameters at five temperature regimes is completely randomized design (CRD). Eventually, the multiple comparison data of averages were done in testing the significant difference whe n they have existed using Tukey\u0026rsquo;s HSD multiple range test (P\u0026thinsp;=\u0026thinsp;05) the calculations were realized by using Spss. For each constant temperatures curves were calculated employing a product limit technique. Population growth rates were computed from the equation of Lotka (23):\u003c/p\u003e \u003cp\u003e1\u0026thinsp;=\u0026thinsp;Σ e-r*x lx * mx (1)\u003c/p\u003e \u003cp\u003eIn which: x\u0026thinsp;=\u0026thinsp;age in days, r\u0026thinsp;=\u0026thinsp;intrinsic rate of increase,\u003c/p\u003e \u003cp\u003eLx\u0026thinsp;=\u0026thinsp;age-specific survival, mx\u0026thinsp;=\u0026thinsp;age-specific number of female offspring. After \"r\" was computed for the original data (rall), differences among rm-values were tested for significance by estimating variances through the jackknife method (24). The jackknife pseudo-value \"rj\" was calculated for the \"n\" samples using the following equation:\u003c/p\u003e \u003cp\u003erj\u0026thinsp;=\u0026thinsp;n * rall - (n-1) *ri (2)\u003c/p\u003e \u003cp\u003eThe mean of \"n\" jackknife pseudo-values for each treatment was subjected to analysis of variance. The scheff\u0026eacute; test was used to compare mean growth rates for different temperature regimes (p\u0026thinsp;=\u0026thinsp;0.05). Because low probability levels were used, there was no concern about inflation of experiment-wise error rates (25). Each of the abovementioned analyses were conducted using the Statgraphics software package.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003eThe developmental time of \u003cem\u003eA. fabae\u003c/em\u003e at five constant temperatures is summarized below (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The results showed that the different instar stages significantly decreased with increasing constant temperatures. The first-instar stage ranged from 1.27 days at 20\u0026deg;C to 3 days at 12\u0026deg;C and there is no significant difference with 1.55 days at 24\u0026deg;C, 1.70 days at 28\u0026deg;C (F\u0026thinsp;=\u0026thinsp;84.03; P\u0026thinsp;=\u0026thinsp;0.05). For the second instar stage there is significant difference between low and high constant temperature ranged from 1.37 days at 28\u0026deg;C to 3.30 days to 12\u0026deg;C (F\u0026thinsp;=\u0026thinsp;63.68; P\u0026thinsp;=\u0026thinsp;0.05). At the third-period from 1.30 days 24\u0026deg;C to 5.80 days at 12\u0026deg;C, but a slight difference at 28\u0026deg;C with 1.75 days (F\u0026thinsp;=\u0026thinsp;96.65; P\u0026thinsp;=\u0026thinsp;0.05). And at fourth instar stage from 1.52 days at 24\u0026deg;C to 4.55 days to 12\u0026deg;C (F\u0026thinsp;=\u0026thinsp;52.73; P\u0026thinsp;=\u0026thinsp;0.05). Moreover, for the total development time (TDT), the same pattern was presented. The instar stage was shorter at high temperature and longer at low temperature. Significant increase was found for TDT at 12\u0026deg;C and 16\u0026deg;C (16.65 and 11.58 days respectively) compared to 20\u0026deg;C, 24\u0026deg;C and 28\u0026deg;C (6.22, 5.70, and 6.62 days respectively) that are statistically similar (Table.1). Concerning the variation days at 28\u0026deg;C treatment. High mortality was observed during immature stages causing a slight difference between the results 24\u0026deg;C and 28\u0026deg;C. The pre-reproduction period showed a significance variation between temperature regimes, increasing at high temperature and increased at low temperature (0.27 days at 28\u0026deg;C to 1.10 days to 16\u0026deg;C F\u0026thinsp;=\u0026thinsp;13.54; P\u0026thinsp;=\u0026thinsp;0.05).\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 Mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SEM) of \u003cem\u003eAphis fabae\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=\"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 \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTemp(\u0026deg;c)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eN1 (day)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eN2 (day)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eN3 (day)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eN4 (day)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eTotal development (day)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePre-reproduction\u003c/p\u003e \u003cp\u003e(day)\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\u003e12\u0026deg;C\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3.30\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e5.80\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4.55\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e16.65\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.97\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1ab\u003c/p\u003e \u003c/td\u003e \u003c/tr\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=\"left\" colname=\"c2\"\u003e \u003cp\u003e2.40\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3.15\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.45\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.70\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e11.58\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.10\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0a\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=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.27\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.57\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.80\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.57\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e6.22\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.32\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0cd\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=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.55\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0cd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.41\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.30\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.52\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e5.70\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.67\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1bc\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=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.70\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.37\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.75\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.80\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e6.62\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.27\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1d\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;d). 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\u003eA linear regression analysis was applied to the developmental level of temperature from 12\u0026deg;C to 28\u0026deg;C range. Within the selected temperature range the developmental rates (r\u003csub\u003e(t)\u003c/sub\u003e) of \u003cem\u003eA. craccivora\u003c/em\u003e increased linearly with the increasing of the temperature (Figure. 2). The theoretical developmental threshold \u003cem\u003eof A craccivora\u003c/em\u003e was recorded as 1.75\u0026deg;C and required 66.66 degree-days for completing all the immature stages to become adult. The number of days of reproduction was significantly longer at low temperature, at 12\u0026deg;C it was 23.95 days (F\u0026thinsp;=\u0026thinsp;31.54: P\u0026thinsp;=\u0026thinsp;0.05) and decreased statistically at higher temperature (28\u0026deg;C; 6.67 days) (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The reproduction period was statistically the same for all temperature regimes and ranged from 0.92 days at 16\u0026deg;C to 2.05 days at 20\u0026deg;C (F\u0026thinsp;=\u0026thinsp;2.09; P\u0026thinsp;=\u0026thinsp;0.05). It was shorter at lower temperature (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Longevity was significantly longer at low temperature (42.33 days at 12\u0026deg;C and 32.6 days at 16\u0026deg;C compared to the others. However, 20\u0026deg;C (22.67 days) was statistically similar to 24\u0026deg;C (20.05 days) and the latter statistically similar to 28\u0026deg;C (16.12 days) (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The life span of \u003cem\u003eA. fabae\u003c/em\u003e days was significantly longer at low temperature 12\u0026deg;C 25.67days and decreased statistically at higher temperature (28\u0026deg;C 9.50 days F\u0026thinsp;=\u0026thinsp;21.50; P\u0026thinsp;=\u0026thinsp;0.05), (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\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\u003eReproduction, post- reproduction, Longevity, life span and number of offspring of \u003cem\u003eAphis fabae\u003c/em\u003e adult female individuals (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SE).\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=\"char\" char=\"\u0026plusmn;\" 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\u003eTemp (\u0026deg;C)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eReproduction (day)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePost- reproduction\u003c/p\u003e \u003cp\u003e(day)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eLongevity (day)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eLife span (day)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eNumber of offspring (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\u003e12\u0026deg;C\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e23.95\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.95\u0026thinsp;\u0026plusmn;\u0026thinsp;0.21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e42.32\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e25.67\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e49.57\u0026thinsp;\u0026plusmn;\u0026thinsp;2.6a\u003c/p\u003e \u003c/td\u003e \u003c/tr\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=\"left\" colname=\"c2\"\u003e \u003cp\u003e18.87\u0026thinsp;\u0026plusmn;\u0026thinsp;1.5b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.92\u0026thinsp;\u0026plusmn;\u0026thinsp;0.24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e32.60\u0026thinsp;\u0026plusmn;\u0026thinsp;1.6b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e21.02\u0026thinsp;\u0026plusmn;\u0026thinsp;1.5ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e62.27\u0026thinsp;\u0026plusmn;\u0026thinsp;5.2a\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=\"left\" colname=\"c2\"\u003e \u003cp\u003e14.00\u0026thinsp;\u0026plusmn;\u0026thinsp;1.3c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e2.05\u0026thinsp;\u0026plusmn;\u0026thinsp;0.45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e22.67\u0026thinsp;\u0026plusmn;\u0026thinsp;1.6c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e16.45\u0026thinsp;\u0026plusmn;\u0026thinsp;1.6bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e55.00\u0026thinsp;\u0026plusmn;\u0026thinsp;4.6a\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=\"left\" colname=\"c2\"\u003e \u003cp\u003e11.47\u0026thinsp;\u0026plusmn;\u0026thinsp;1.1c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e1.42\u0026thinsp;\u0026plusmn;\u0026thinsp;0.47\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e20.05\u0026thinsp;\u0026plusmn;\u0026thinsp;1.3cd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e14.35\u0026thinsp;\u0026plusmn;\u0026thinsp;1.3cd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e48.67\u0026thinsp;\u0026plusmn;\u0026thinsp;4.5a\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=\"left\" colname=\"c2\"\u003e \u003cp\u003e6.67\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e2.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e16.12\u0026thinsp;\u0026plusmn;\u0026thinsp;1.2d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e9.50\u0026thinsp;\u0026plusmn;\u0026thinsp;1.2d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e16.72\u0026thinsp;\u0026plusmn;\u0026thinsp;2.5b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"6\"\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;d). 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\u003eVirtually at the high level of temperatures, the mother aphids were less survived. The number of newborns per days, their size was reduced and early died at 28\u0026deg;C (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). This study was resulted that with an increasing of constant temperatures, death ratio of the immature populations was more effectively. For example, at 28\u0026deg;C, there is 25% of death compared to the lowest temperature regimes The number of generation are produced per (To) \u003cem\u003eA. fabae\u003c/em\u003e at 12\u0026deg;C were 29.14 generations while at 28\u0026deg;C 11.50 generations (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The net reproduction (Ro) was higher at 16\u0026deg;C (62.27 aphids/aphid) while at low at 28\u0026deg;C (16.72 aphids/aphid) (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The offspring per reproduction days was significantly longer at 24\u0026deg;C 4.84 aphids/days compared to others. \u003cem\u003eA. fabae\u003c/em\u003e at middle temperatures was well developed and showed the highest population growth per capita rate across the intrinsic rate of increase ranged at 20\u0026deg;C and 24\u0026deg;C (0.35 and 0.36 aphids/aphid per days) and the lowest at 12\u0026deg;C (0.15 aphids/aphid per day). The number of offspring was significantly different at 28\u0026deg;C 16. 72 aphids compared to the others.\u003c/p\u003e \u003cp\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\u003eGeneration time (T0), net reproduction (Ro), and intrinsic rate of increase (rm) of \u003cem\u003eAphis fabae\u003c/em\u003e on bean leaf discs at five temperatures levels\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"8\"\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=\".\" 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=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\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\u003eSample size(N)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFemale adults(N)\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\u003eGeneration time(T\u003csub\u003e0\u003c/sub\u003e)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eReproduction rate (R\u003csub\u003e0\u003c/sub\u003e)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eOffspring per production day\u003c/p\u003e \u003cp\u003e(Mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SEM)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eIntrinsic rate\u003c/p\u003e \u003cp\u003eincrease(aphis/aphid/d)\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\u003e12\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=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e29.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e50.96\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.7b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.15\u003c/p\u003e \u003c/td\u003e \u003c/tr\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=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e23.17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e62.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.21\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=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e38\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\u003e14.40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e55\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e3.99\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2 ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.35\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=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e38\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\u003e12.97\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e48.68\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e4.84\u0026thinsp;\u0026plusmn;\u0026thinsp;0.5a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.36\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=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e11.50\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e16.72\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0.27\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"8\"\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;d). 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"},{"header":"Discussion","content":"\u003cp\u003eTemperature affects the development as well as growth time; consequently, the evolution and dynamics of insect populations in the field are dictated by ambient temperature. For this reason, there is a considerable interest for a long time in the temperature relationships of development time, reproduction, nymphal mortality rate, longevity, and fecundity of A. \u003cem\u003efabae.\u003c/em\u003e According to the outcomes of this study transparently, showed that at five constant temperatures on biological development of A. \u003cem\u003efaba\u003c/em\u003ee (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, \u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e, \u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The different nymphal stages and total development significantly decreased from 12\u0026deg;C until 24\u0026deg;C with increasing constant temperature, however from 24\u0026deg;C to 28\u0026deg;C of the total developmental was augmented again. According to some similar studies were reported by (Tofangsazi et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Akca et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Insect cultured at the temperatures over the threshold develop more slowly than others that reared below favorable conditions (Satar et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e1999\u003c/span\u003e and \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Kersting et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Dona and Satar (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Another observation has been related by Olmez et al., 2003; Maelzer, 1977) who reported that the developmental rate of M. rosae was retarded at temperatures above 25\u0026deg;C to 32\u0026deg;C. This study presented the same reality for A. \u003cem\u003efabae\u003c/em\u003e on faba bean at 28\u0026deg;C showing a significant increase in developmental time evaluated with that at 24\u0026deg;C. Therefore, only at 20\u0026deg;C in the second-instar stage, a slight increase has been presented comparing with 24\u0026deg;C. The percentage of mortality at 28\u0026deg;C has been shown highest elevation death, while at 12\u0026deg;C, the smallest it resists at hundred percent (100%). According to the physiology of A. \u003cem\u003efabae\u003c/em\u003e, its adaptation reputes more for cold temperatures than high. Although it occurs in tropical areas of Africa, it is not a very serious pest in warm environments (Farhadi et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Generally, aphids at lower temperatures need to take more developmental time before they start to produce nymphs (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). reproduction period, the duration of reproduction of \u003cem\u003eA. fabae\u003c/em\u003e clearly showed as much as decreasing over threshold temperature limit take more time for reproducing. For example, at 12\u0026deg;C 23.95 days to 28\u0026deg;C 6.67days. The post- reproduction period didn\u0026rsquo;t show any significant differences between the temperatures (Table.2) Longevity and life span from some previous studies reported that decreased constant temperature engenders an increasing period. It\u0026rsquo;s a common phenomenon from various aphid species (Morgan et al., \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Wang \u0026amp; Tsai, 2000; Collins \u0026amp; Leather, 2001; Satar \u0026amp; Yokomi, 2002). Another observation has been related by (Olmez et al., 2003; Akca et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2015\u003c/span\u003e and Maelzer. 1977) who reported that the developmental rate of M. \u003cem\u003erosae\u003c/em\u003e was retarded at temperatures above 25\u0026deg;C. While in our study over 24\u0026deg;C similar results were presented.\u003c/p\u003e \u003cp\u003eThe intrinsic rate of natural increase (\u003cem\u003er\u003c/em\u003e) is a key an indicator that can be demonstrated as the most favorable temperature level for better population growth because it comes from all over impacts caused by the temperature on the life table of a population. The population reared from 12 to 24\u0026deg;C, this last presented the highest (\u003cem\u003er-\u003c/em\u003e value compare with others (\u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.36 aphids/aphid per day) because it the most rapid development, the highest survival of nymphal stages in an average of number 4.80 nymphs of daily progeny and proved as long as \u003cem\u003er\u003c/em\u003e -value exposed at an alternate temperature couldn\u0026rsquo;t be subjected as the highest net production rate per female for A. \u003cem\u003efabae\u003c/em\u003e, but in this case with an increasing of constant temperature 24\u0026deg;C to 16\u0026deg;C (\u003cem\u003eRo\u003c/em\u003e\u0026thinsp;=\u0026thinsp;48.68 to 62.27). A similar result has been reported by (Akca et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) gradually decreasing from 15 to 25\u0026deg;C (\u003cem\u003er\u0026thinsp;=\u003c/em\u003e\u0026thinsp;0.43 d-1) and the opposite for (\u003cem\u003eRo\u003c/em\u003e\u0026thinsp;=\u0026thinsp;90.44 to 77.46) increased. According to those results from the smallest to the biggest the most suitable temperature of A, \u003cem\u003efabae\u003c/em\u003e ranged 16 to 24\u0026deg;C, otherwise, its biology can be affected greatly. On the other hand, the host variety also can play a major role in aphid biology. Related by (Fernandez-Quintanilla et al., 2002) about a range of life parameters of A. \u003cem\u003efabae\u003c/em\u003e on weed species in winter and summer as 0.14 to 0.36 days (the mean unit used in this article was nymphs per female per days).\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003eConflict of Interest The authors declare that they have no conflict 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:\u003c/strong\u003e We would like to thank the team management of the Project Development and Coordination Unit. This research was part of the Msc of Rochelyn DONA was funded by Project Development and Coordination Unit from \u0026Ccedil;ukurova University (Grants Code: FYL-2020-12688).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding:\u0026nbsp;\u003c/strong\u003eThe study was supported by Cukurova University. (Grants Code: FYL-2020-12688).\u003c/p\u003e\n\u003cp\u003eAuthors \u0026lsquo;contributions RD and SS conceived, designed and performed research. Both of the authors analyzed data, and wrote the maniscript. Both authors read and approved the manuscript.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAdabi, S. T., Talebi, A. A., Fathipour, Y., \u0026amp; Zamani, A. A. (2010). Life history and demographic parameters of Aphis fabae (Hemiptera: Aphididae) and its parasitoid, Aphidius matricariae (Hymenoptera: Aphidiidae) on four sugar beet cultivars. \u003cem\u003eActa entomologica serbica\u003c/em\u003e, \u003cem\u003e15\u003c/em\u003e(1), 61-73.\u003c/li\u003e\n\u003cli\u003eAkca, I., Ayvaz, T., Yazici, E., Smith, C. L., \u0026amp; Chi, H. (2015). Demography and population projection of Aphis fabae (Hemiptera: Aphididae): with additional comments on life table research criteria. \u003cem\u003eJournal of Economic Entomology\u003c/em\u003e, \u003cem\u003e108\u003c/em\u003e(4), 1466-1478.\u003c/li\u003e\n\u003cli\u003eAl-Antary, T., Ateyyat, M., \u0026amp; Thaliji, T. (2007). Effect of eight cultivars of faba bean, Vicia faba, on the black bean aphid, Aphis fabae Scopoli (Homoptera: Aphididae) in the field. \u003cem\u003eDirasat (Agric. 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Laboratory studies on aggregation, size and fecundity in the black bean aphid, Aphis fabae Scop. \u003cem\u003eBulletin of Entomological Research\u003c/em\u003e, \u003cem\u003e61\u003c/em\u003e(1), 97-111.\u003c/li\u003e\n\u003cli\u003eDona, R., \u0026amp; Satar, S. (2024). Thermal effects on the biological parameters of the bean \u003cem\u003eAphis craccivora\u003c/em\u003e (Hemiptera: Aphididae).\u003c/li\u003e\n\u003cli\u003eEsmaeili-Vardanjani, M., Askarianzadeh, A., Saeidi, Z., Hasanshahi, G. H., Karimi, J., \u0026amp; Nourbakhsh, S. H. (2013). A study on common bean cultivars to identify sources of resistance against the black bean aphid, Aphis fabae Scopoli (Hemiptera: Aphididae). \u003cem\u003eArchives of Phytopathology and Plant Protection\u003c/em\u003e, \u003cem\u003e46\u003c/em\u003e(13), 1598-1608.\u003c/li\u003e\n\u003cli\u003eEsmaeili-Vardanjani, M., Askarianzadeh, A., Saeidi, Z., Hasanshahi, G. H., Karimi, J., \u0026amp; Nourbakhsh, S. H. (2013). 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Identification of sources of resistance to the black bean aphid, Aphis fabae Scopoli, in Faba bean, Vicia faba L., accessions. \u003cem\u003eJournal of Crop Protection\u003c/em\u003e, \u003cem\u003e4\u003c/em\u003e(2), 217-224.\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"international-journal-of-tropical-insect-science","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jtis","sideBox":"Learn more about [International Journal of Tropical Insect Science](http://link.springer.com/journal/42690)","snPcode":"42690","submissionUrl":"https://www.editorialmanager.com/jtis/default2.aspx","title":"International Journal of Tropical Insect Science","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Developmental time, intrinsic rate, optimal range, reproduction period, temperature regimes","lastPublishedDoi":"10.21203/rs.3.rs-5220155/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-5220155/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe aim of this study was to evaluate the thermal effects on the biological parameters of \u003cem\u003eAphis fabae\u003c/em\u003e Scopoli (Hemiptera: Aphididae). The developmental times, survival, and reproductive data were collected for \u003cem\u003eAphis fabae\u003c/em\u003e reared on detached bean leaves (\u003cem\u003ePhaseolus vulgaris\u003c/em\u003e L.\u003cem\u003e)\u003c/em\u003e \u0026lsquo;pinto beans at five temperatures regimes (12\u0026deg;C, 16\u0026deg;C, 20\u0026deg;C, 24\u0026deg;C and 28\u0026deg;C), 65% relative humidity (RH), and a photoperiod of 16:8 (L: D) h. The developmental times of immature stages ranged from 16,65 days at 12\u0026deg;C to 5.70 days at 24\u0026deg;C, but a slight increase again at 28\u0026deg;C (6.62 days). This study presented the optimal developmental threshold for A. \u003cem\u003efabae\u003c/em\u003e slightly to 24\u0026deg;C. The average longevity of mature females significantly decreased from 42.32 days at 12\u0026deg;C to 16.12 days at 28\u0026deg;C. The reproduction rate per female was 62.27 individuals at 16\u0026deg;C and 12.72 individuals at 28\u0026deg;C. The mean generation period of the population ranged from 29.24 at 12\u0026deg;C to 11.50 at 28\u0026deg;C. The highest intrinsic rate of increase (\u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.41) was recorded at 24\u0026deg;C, the lowest at 12\u0026deg;C (\u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.15). It was evident that temperatures over 28\u0026deg;C augmented the development time, accelerated the death ratio of the nymphal stages, Shrunk Adult longevity, and reduced fecundity. The optimal range of temperature for population growth of \u003cem\u003eA. fabae\u003c/em\u003e on bean was 16\u0026deg;C-24\u0026deg;C according to this study.\u003c/p\u003e","manuscriptTitle":"Thermal effects on the growth and developmental time of bean Aphis fabae (Hemiptera: Aphididae)","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-05-14 17:03:03","doi":"10.21203/rs.3.rs-5220155/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Major revisions","date":"2025-06-03T08:33:06+00:00","index":"","fulltext":""},{"type":"reviewerAgreed","content":"","date":"2025-05-14T05:33:16+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-05-09T12:05:11+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-10-17T05:32:05+00:00","index":"","fulltext":""},{"type":"submitted","content":"International Journal of Tropical Insect Science","date":"2024-10-15T18:43:13+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"international-journal-of-tropical-insect-science","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jtis","sideBox":"Learn more about [International Journal of Tropical Insect Science](http://link.springer.com/journal/42690)","snPcode":"42690","submissionUrl":"https://www.editorialmanager.com/jtis/default2.aspx","title":"International Journal of Tropical Insect Science","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"b7f711c4-255b-4626-bedb-aff39ae78255","owner":[],"postedDate":"May 14th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2025-11-05T08:53:35+00:00","versionOfRecord":[],"versionCreatedAt":"2025-05-14 17:03:03","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-5220155","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-5220155","identity":"rs-5220155","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}

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