Hemoglobin Genotypes in Turbot (Scophthalmus maximus Rafinesque), Their Oxygen Affinity Properties and Relation With Growth

A growth trial lasting 15 months was conducted with 80 individually tagged turbot at four temperatures (10, 13, 16, and 19°C). Haemoglobins of the individually tagged fish were analysed by agar gel electrophoresis and the fish segregated into three genotypes named; Hb-I(1/1), Hb-I(1/2) and Hb-I(2/2). The genotype specific growth rates of the fish were analysed. In addition, the oxygen affinity properties of the hemoglobin genotypes were studied at 10 and 19°C. The genotype Hb-I(2/2) displayed the overall best growth at 10, 13 and 16°C, but no differences in growth among genotypes were seen at 19°C. Further, Hb-I(2/2) had the highest oxygen affinity at 10 and 19°C, whereas Hb-I(1/1) had the lowest oxygen affinity at both temperatures. In conclusion, the hemoglobin polymorphism in turbot seems to be correlated with physiological performance, and thus useful in future selection programs for turbot as a correlated trait with production characteristics.     

Genetic variation and functional properties of Atlantic cod hemoglobins: introducing a modified tonometric method for studying fragile hemoglobins

Hemoglobin polymorphism in Atlantic cod has been investigated with respect to physiological performance at 10, 15 and 20 degrees C applying a modified tonometric method for O2 equilibrium analysis with full control of the equilibrating gas mixture. The results did not indicate any dissociation of the hemoglobins by a reduction in cooperativity and a parallel increase in affinity during the analytical procedure in contrast to the original tonometric method. With the applied preparation technique, we could store the hemolysate for 70 days at -25 degrees C without any significant changes in the O2 binding properties (P < 0.05) demonstrating the high quality of this procedure for analysing fragile fish hemoglobins. The present investigation demonstrates that the oxygen affinity of the hemoglobins varied between the genotypes. At all temperatures, except 20 degrees C and pH 8.0, Hb-I(2/2) had a higher O2 affinity than Hb-I(1/1). These results conform with previous results (16), suggesting Hb-I(2/2), the genotype which is the dominant allele in northern areas, to be the most efficient O2 carrier at low temperatures. The highest O2 affinity, however, was found for Hb-I(2/2b), supporting the results of Fyhn et al. (9), that this genotype is more restricted to coastal and warmer water and thus a better marker of the coastal population. Our results further suggest a correlation between genotype specific growth rates and oxygen affinities at all temperatures studied, with the highest growth rates observed in those genotypes having the highest O2 affinities. In conclusion, the hemoglobin polymorphism of cod seems to be correlated with physiological performance.     

Temperature acclimation modulates the oxygen binding properties of the Atlantic cod (Gadus morhua L.) genotypes-HbI*1/1, HbI*1/2, and HbI*2/2-by changing the concentrations of their major hemoglobin components (results from growth studies at different temperatures)

The influence of long-term acclimation temperatures in Atlantic cod (Gadus morhua) was studied by growth experiments carried out over a total of 272 individuals. The attention focused on the structural and functional modulation of the five electrophoretically distinguishable genotypes of cod hemoglobin (HbI*1/1, HbI*1/2, HbI*2/2, HbI*1/2b, and HbI*2/2b) and on the correlation with body length/weight. The main results can be summarized as follows. (1) Acclimation to lower (4 and 8 degrees C) and higher (12 and 15 degrees C) temperatures favors the expression of, respectively, more anodic and more cathodic hemoglobin components. (2) The optimal O(2) transporting features are observed at 12 degrees C, as well as a saturation-dependent temperature dependence of O(2) binding, which furthermore is strongly dependent upon the acclimation background. (3) The optimal growth condition for the three main genotypes (HbI*1/1, HbI*1/2, and HbI*2/2) is associated with T=12 degrees C. The overall results are consistent with the idea that environmental temperatures constitute a primary factor in the aggregation of individuals physiologically more than genetically homogeneous. This is fully confirmed by careful statistical analysis carried out over a subset of individuals for which the full set of structural (isoelectric focusing), functional (O(2) binding), and growth data was available.     

Hypoxic avoidance behaviour in cod (Gadus morhua L.): The effect of temperature and haemoglobin genotype

Hypoxia can influence fish growth, survival and on larger scales, population structure. These effects may be influenced by water temperature, and may vary intra-specifically with genotype. In Atlantic cod (Gadus morhua L.), the two haemoglobin homozygotes (Hb-I?11 and Hb-I?22) vary in oxygen affinity at different temperatures, which is thought to correspond to variation in hypoxia tolerance. We therefore tested if hypoxic avoidance behaviour in cod 1) depends on ambient temperature and 2) is modified by haemoglobin genotype. In a laminar flow choice box, we subjected juvenile cod to an initial phase of non-escapable hypoxia, and a subsequent recovery phase, where one habitat was kept at 20% O₂ saturation while the other was raised in steps to full saturation. The experiment was performed at 5 and 15 °C with Hb-I?11 and Hb-I?22 cod. Cod responded to inescapable hypoxia by reducing their overall swimming speed and then, at the initial levels of the recovery phase, avoiding the most hypoxic habitat, irrespective of temperature or genotype. Fish recovered quickly as O₂ levels increased, as evidenced by increased swimming speed and time spent in the most hypoxic habitat. The avoidance response depended strongly on temperature: the relative reduction in speed and avoidance of the most hypoxic habitat was more pronounced at 15 than at 5 °C. During the recovery phase, stressed fish initially maintained a higher swimming speed in the most hypoxic habitat. However, as O₂ increased, swimming speed in both habitats converged. This point of convergence occurred at a lower O₂ saturation at 5 °C. Fish ventilation rate in inescapable hypoxia was also higher at 15 °C. Haemoglobin genotype did not influence either ventilation rates or the nature of the hypoxic avoidance response at either temperature, but Hb-I?11 cod swam faster than Hb-I?22 cod in normoxia at 15 °C. Our results indicate that increased temperature limits the ability of cod of both haemoglobin genotypes to exploit hypoxic habitats. This may have negative future consequences for coastal cod stocks in light of increasing global temperatures and eutrophication in coastal waters.     

Effect of precocene II on fatty acid metabolism in the pea aphid, Acyrthosiphon pisum, under cold stress

Pea aphids, Acyrthosiphon pisum (Harris), reared at 10 degrees C contain higher levels of fatty acids than those reared at 25 degrees C. This is primarily the result of an accumulation of triacylglycerols containing myristic acid. When aphids reared at 25 degrees C were transferred to 10 degrees C there was a gradual increase in triacylglycerol content that reached a maximum at 16 days post-transfer. Treatment of aphids with precocene II prior to transfer to 10 degrees C blocked the accumulation of fatty acids including myristic acid. A single application of 2 microg precocene II/aphid or two applications of 0.5 microg precocene II/ aphid administered on consecutive days resulted in aphids moved to 10 degrees C maintaining the same fatty acid profile as aphids maintained at 25 degrees C. Aphids that were treated with precocene II and maintained at 25 degrees C did not show changes in fatty acid profiles. Rearing aphids at 10 degrees C resulted in lower rates of reproduction and lower total numbers of progeny with longer longevity. Treatment with precocene II significantly decreased the total number of progeny produced at both temperatures. Precocene II did not reduce life span of aphids reared at 25 degrees C, however, the life span of treated aphids reared at 10 degrees C was decreased. The mechanism by which precocene II prevents the accumulation of myristic acid in aphids reared at 10 degrees C remains to be determined.     

Influence of temperature on growth rate and competition between two psychrotolerant Antarctic bacteria: low temperature diminishes affinity for substrate uptake.

The growth kinetics of two psychrotolerant Antarctic bacteria, Hydrogenophaga pseudoflava CR3/2/10 (2/10) and Brevibacterium sp. strain CR3/1/15 (1/15), were examined over a range of temperatures in both batch culture and glycerol-limited chemostat cultures. The maximum specific growth rate (mu max) and Ks values for both bacteria were functions of temperature, although the cell yields were relatively constant with respect to temperature. The mu max values of both strains increased up to an optimum temperature, 24 degrees C for 2/10 and 20 degrees C for 1/15. Strain 1/15 might therefore be considered to be more psychrophilic than strain 2/10. For both bacteria, the specific affinity (mu max/Ks) for glycerol uptake was lower at 2 than at 16 degrees C, indicating a greater tendency to substrate limitation at low temperature. As the temperature increased from 2 to 16 degrees C, the specific affinity of 1/15 for glycerol increased more rapidly than it did for 2/10. Thus 1/15, on the basis of this criterion, was less psychrophilic than was 2/10. The steady-state growth kinetics of the two strains at 2 and 16 degrees C imply that 1/15 would be able to outgrow 2/10 only at relatively low substrate concentrations (< 0.32 g of glycerol.liter-1) and high temperatures (> 12 degrees C), which suggests that 1/15 has a less psychrotolerant survival strategy than does 2/10. Our data were compared with other data in the literature for bacteria growing at low temperatures. They also showed an increase of substrate-specific affinity with increasing temperature.(ABSTRACT TRUNCATED AT 250 WORDS)     

Black soldier fly (Diptera: Stratiomyidae) larval heat generation and management

Mass production of black soldier fly, Hermetia illucens (L.) (Diptera: Stratiomyidae), larvae results in massive heat generation, which impacts facility management, waste conversion, and larval production. We tested daily substrate temperatures with different population densities (i.e., 0, 500, 1000, 5000, and 10 000 larvae/pan), different population sizes (i.e., 166, 1000, and 10 000 larvae at a fixed feed ratio) and air temperatures (i.e., 20 and 30 °C) on various production parameters. Impacts of shifting larvae from 30 to 20 °C on either day 9 or 11 were also determined. Larval activity increased substrate temperatures significantly (i.e., at least 10 °C above air temperatures). Low air temperature favored growth with the higher population sizes while high temperature favored growth with low population sizes. The greatest average individual larval weights (e.g., 0.126 and 0.124 g) and feed conversion ratios (e.g., 1.92 and 2.08 g/g) were recorded for either 10 000 larvae reared at 20 °C or 100 larvae reared at 30 °C. Shifting temperatures from high (30 °C) to low (20 °C) in between (∼10-d-old larvae) impacted larval production weights (16% increases) and feed conversion ratios (increased 14%). Facilities should consider the impact of larval density, population size, and air temperature during black soldier fly mass production as these factors impact overall larval production.     

Laboratory environment effects on the reproduction and mortality of adult screwworm (Diptera: Calliphoridae)

The New World screwworm, Cochliomyia hominivorax Coquerel, is mass reared for screwworm eradication initiatives that use the sterile insect technique. New methods for rearing have helped to reduce the cost of the eradication program. We examined the effect and interaction of three temperatures (24.5, 29.5 and 34.5 degrees C), two diets (2% spray-dried blood plus 0.05% vitamins and corn syrup carrageenan) and three population densities (300, 400, and 500 flies/cage) on egg production, egg hatch, number of observable fertilized eggs, mortality (male and female) and ovarian development. The three population densities did not affect any of the parameters monitored. Using the protein diet increased egg production at all temperatures. Diet did not affect egg hatch or female mortality. Male mortality was significantly greater when fed the protein diet and reared at 24.5 degrees C and 34.5 degrees C. Egg hatch was significantly less when the flies were reared at 34.5 degrees C. When exposed to high temperatures (37 degrees C and 40 degrees C) egg production, egg hatch, fertility and mortality were adversely affected. At the higher temperatures, yolk did not adequately form during oogenesis. When compared to the normal rearing photoperiod (12 L:12 D), short photoperiod (1 L:23 D) increased egg production, egg hatch and fertility but lowered mortality.     

Observations on growth rates of Arctic charr, Salvelinus alpinus (L.), reared at low temperature

Food intake and growth of Arctic charr, Salvelinus alpinus , in fresh water was studied at three temperatures 2·9, 8·4 and 13·1° C). Best growth and highest food intake occurred at 13·1°C. The approximate chemical composition was dependent upon rearing temperature, fish reared at the highest temperature depositing large quantities of fat. Fish were later grown on in either fresh or salt water where two growth patterns were observed. In the light of published data for Salvelinus spp., it is suggested that the poorest growth was shown by fish which were incapable of complete adaptation to saltwater conditions.     

Effect of different thermal conditions on the pre-imaginal biology of Culex apicinus (Philippi, 1865) (Diptera: Culicidae).

The time of development and survival of the pre-imaginal period and the adult body size of Culex apicinus were analyzed in individuals reared from the 1st instar larva to adult emergence under laboratory and field conditions. In the laboratory, insects were exposed to three constant temperatures (15, 20, and 25 degrees C) and a photoperiod of 14:10 (L:D). In the field, temperature and photoperiod were not manipulated; during the study period water temperature ranged between 15.5 and 24.2 degrees C, and photoperiod changed from 13:11 to 14:10 (L:D). Survival to the imaginal stage at 15 degrees C (85.4%) and in the field (88.8%) was higher than that at 20 degrees C (45.8%) and 25 degrees C (8.3%). Based on femur and wing length measurements, the mosquitoes reared under field conditions (mean water temperature = 20.5 degrees C) were smaller than those laboratory-reared, even when the time of development in the field (29.5 - 36 days) was longer than that observed at 15 degrees C (24 days), 20 degrees C (15 days), and 25 degrees C (11 days). Male emergence was earlier than that of females (protandry) only in the field. Although laboratory conditions included a photoperiod and temperature range comparable with the observed values in the field, the disagreements between field and laboratory results suggest that the characteristics examined in this work could be affected by the joint variation of several factors not controlled in field conditions, such as photoperiod, temperature regime, and/or food quality of larval habitat.     

Effects of Temperature and Photoperiod on Flowering in Lentils (Lens culinaris Medic.)

Factorial combinations of three photoperiods (10, 13 and 16h), two day temperatures (18 and 28 C) and two night temperatures(5 and 13 C) were imposed on nodulated plants of six diversegenotypes (cultivars and land-races) of lentil (Lens culinarisMedic.) grown in pots in growth cabinets from vernalized (1.50.5 C for 30d) or non-vernalized seeds (i.e. 144 ‘treatment’combinations). The times from sowing to the appearance of firstopen flowers were recorded. Vernalization, long days and warmtemperatures hastened flowering but genotypes differed in relativesensitivity to each of these factors and in time to floweringin the same most-inductive environment. Rates of progress towardsflowering (i.e. 1/f the reciprocals of the times to first flower,f) in all genotypes, vernalized or not, were linear functionsof both mean temperature,     

Temperature until the 'eyed stage' of embryogenesis programmes the growth trajectory and muscle phenotype of adult Atlantic salmon

We investigated how adult growth in Atlantic salmon (Salmo salar L.) was affected by changing embryonic temperature from fertilization until the completion of eye pigmentation. Fertilized eggs from several hundred families were divided between four temperature treatments (2, 5, 8 or 10 degrees C) and subsequently reared in identical conditions in replicated tanks. Fish exposed to 2 and 5 degrees C treatments were significantly smaller at smoltification than groups at higher temperatures, but showed substantial compensatory catch-up growth. Remarkably, temperature during this short window of embryogenesis dictated adult myogenic phenotype three years later with significant treatment effects on the muscle fibre final number (FFN), maximum diameter, nuclear density and size distribution. FFN was highest for the 5 degrees C treatment and was reduced at higher and lower treatment temperatures. Our results require direct temperature effects on embryonic tissues, such as the stem cell-containing external cell layer, in order to produce persistent effects on juvenile and adult growth.     

Rearing temperature influences flavivirus vector competence of mosquitoes

Culex annulirostris Skuse mosquitoes (Brisbane strain) were reared at 20 degrees C or 27 degrees C and the adult females were experimentally infected by feeding Murray Valley encephalitis virus (MVE). They were then maintained (a) in the insectary at 20 degrees C, after rearing at either 20 degrees C or 27 degrees C; (b) at ambient outdoor temperatures, range 12.2-28.9 degrees C, mean 19.6 degrees C; or (c) at 27 degrees C after rearing at 27 degrees C. There was no significant difference in rates of MVE infection or transmission when mosquitoes were reared and maintained constantly at 20 degrees C or 27 degrees C. However, for females kept at reduced temperature (i.e. mean = 19.6 degrees C or 20 degrees C after rearing at 27 degrees C), the infection and transmission rates of MVE were significantly reduced (2 x 8 replicates). This investigation illustrates that vector competence is depressed by decreasing temperatures for adult mosquitoes compared with those they experienced during development. Similar patterns were evident with previously published work on Japanese and St Louis encephalitis, dengue and yellow fever. The process appears to be reversible, i.e. increased temperature raises virus infection and transmission rates. It is concluded that, without incubation at warmer temperatures, flavivirus recovery from overwintering mosquitoes will be negatively biased.     

Interactive effects of rearing temperature and oxygen on the development of Drosophila melanogaster

Although higher temperatures strongly stimulate ectothermic metabolic rates, they only slightly increase oxygen diffusion rates and decrease oxygen solubility. Consequently, we predicted that insect gas exchange systems would have more difficulty meeting tissue oxygen demands at higher temperatures. In this study, Drosophila melanogaster were reared from egg to adult in hyperoxic (40%), hypoxic (10%), and normoxic (21%) conditions and in temperatures ranging from 15 degrees -31.5 degrees C to examine the interactive effect of temperature and oxygen on development. Hyperoxia generally increased mass and growth rate at higher rearing temperatures. At lower rearing temperatures, however, hyperoxia had a very small effect on mass, did not affect growth rate, and lengthened time to eclosion. Relative to normoxia, flies reared in hypoxic conditions were generally smaller (mass and thorax length), had longer eclosion times, slower growth rates, and reduced survival. At cooler temperatures, hypoxia had relatively modest or nonsignificant effects on development, while at higher temperatures, the effects of hypoxia were large. These results suggest that higher temperatures reduce oxygen delivery capacity relative to tissue oxygen needs, which may partially explain why ectotherms are smaller when development occurs at higher temperatures.     

Development, survivorship, and reproduction of Helicoverpa armigera (Lepidoptera: Noctuidae) under constant and alternating temperatures

Laboratory studies were conducted to assess the effect of temperature on the survival, development, fecundity, and longevity of Helicoverpa armigera (Hübner) at 11 constant temperatures ranging from 12.5 to 40 degrees C, as well as at five alternating temperature regimes (25-10, 30-15, 32.5-17.5, 35-20, and 35-27.5 degrees C) and under a photoperiod of 16:8 (L:D) h. H. armigera reared at constant temperatures did not develop from egg to adult (emergence) outside the temperature range of 17.5-32.5 degrees C. The alternating conditions expanded this range from 10 to 35 degrees C. The lowest developmental thresholds of the immature stages were estimated by a linear model and ranged from 10.17 (pupal stage) to 11.95 degrees C (egg stage) at constant temperature regimes and from 1.1 to 5.5 degrees C, respectively at alternating temperatures. The values of developmental thresholds estimated using the nonlinear (Lactin-2) model were lower than those estimated by the linear model for constant and alternating temperature regimes except for larval and pupal stages at constant temperatures. Mean adult longevity fluctuated from 34.4 d at 15 degrees C to 7.6 d at 35 degrees C. Females reared under all alternating temperature regimes laid more eggs than females reared at any, except the 25 degrees C, constant temperature treatment. The intrinsic rate of increase was highest at 27.5 degrees C, at both the constant and the corresponding alternating temperature regimes (0.147 and 0.139, respectively). Extreme temperatures had a negative effect on life table parameters.     

Respiration as a percentage of daily photosynthesis in whole plants is homeostatic at moderate, but not high, growth temperatures

Here, we investigated the impact of temperature on the carbon economy of two Plantago species from contrasting habitats. The lowland Plantago major and the alpine Plantago euryphylla were grown hydroponically at three constant temperatures: 13, 20 and 27 degrees C. Rates of photosynthetic CO(2) uptake (P) and respiratory CO(2) release (R) in shoots and R in roots were measured at the growth temperature using intact plants. At each growth temperature, air temperatures were changed to establish short-term temperature effects on the ratio of R to P (R/P). In both species, R/P was essentially constant in plants grown at 13 and 20 degrees C. However, R/P was substantially greater in 27 degrees C-grown plants, particularly in P. euryphylla. The increase in R/P at 27 degrees C would have been even greater had biomass allocation to roots not decreased with increasing growth temperature. Short-term increases in air temperature increased R/P in both species, with the effects of air temperature being most pronounced in 13 degrees C-grown plants. We conclude that temperature-mediated changes in biomass allocation play an important role in determining whole-plant R/P values, and, while homeostasis of R/P is achieved across moderate growth temperatures, homeostasis is not maintained when plants are exposed to growth temperatures higher than usually experienced in the natural habitat.     

Purification, crystallisation and preliminary X-ray study of haemoglobin from Crocodilis palustris and Crocodilis porosus

The unsolved three-dimensional structure of crocodile haemoglobin and its prospects as a blood substitute have led us to initiate the purification and crystallisation of haemoglobin molecules from crocodile species (Crocodilis palustris or mugger and Crocodilis porosus or salt water crocodile). The work has resulted in the prevention of polymerisation of naked haemoglobin molecules using N-ethylmaleimide or iodoacetamide. The purified monomeric haemoglobin molecule of C. porosus was crystallised in two different forms and X-ray diffraction data were collected up to 2 A resolution for both forms. Form I: a=53.62, b=53.55, c=103.77 A; beta=93.35 degrees, space group P2(1), Z=2. Form II: a=71.30, b=54.70, c=80.00 A; beta=106.4 degrees, space group P2(1), Z=2. Structure solution and rigid body refinement of form I data resulted in a model with R(free)=0.42 and R=0.35.     

The immune response of tilapia Oreochromis mossambicus and its susceptibility to Streptococcus iniae under stress in low and high temperatures

Mozambique tilapia Oreochromis mossambicus acclimated to 27 degrees C were then held at 19, 23, 27 (control), 31 and 35 degrees C, and were examined for non-specific cellular and humoral responses after 12-96 h. Total leucocyte count decreased significantly when fish were transferred to 19 and 23 degrees C after 48 and 96 h, and when transferred to 35 degrees C over 12-96 h, respectively. Respiratory burst decreased significantly when fish were transferred to 19, 31 and 35 degrees C over 24-96 h, whereas phagocytic activity and phagocytic index decreased significantly when fish were transferred to low temperatures (19 and 23 degrees C) and high temperatures (31 and 35 degrees C) over 12-96 h. Lysozyme activity decreased significantly when fish were transferred to 19 degrees C after 12-96 h, but increased significantly when transferred to 31 and 35 degrees C over 48-96 h. Alternative complement pathway (ACH(50)) also decreased significantly when transferred to 19 and 23 degrees C after 12h, but increased significantly when transferred to 31 and 35 degrees C after 24h. In another experiment, tilapia reared at 27 degrees C were injected intraperitoneally with Streptococcus iniae at a dose of 1 x 10(7)colony-forming units (cfu)fish(-1), and then reared onward at water temperatures of 19, 23, 27 (control), 31 and 35 degrees C. Over 48-168 h, the cumulative mortality of S. iniae-injected fish held in 19 and 35 degrees C was significantly higher than that of injected-fish held in 23, 27 and 31 degrees C. It is concluded that transfer of tilapia O. mossambicus from 27 degrees C to low temperatures (19 and 23 degrees C) after 12h, and transfer of fish from 27 degrees C to high temperatures (31 and 35 degrees C) reduced their immune capability. Moreover, tilapia under temperature stress at 19 and 35 degrees C from 27 degrees C decreased its resistance against S. iniae.     

A true summer diapause induced by high temperatures in the cotton bollworm, Helicoverpa armigera (Lepidoptera: Noctuidae)

Summer diapause in the cotton bollworm, Helicoverpa armigera (Hübner), which prolongs the pupal stage, particularly in males, is induced by high temperatures. In the laboratory, summer-diapausing pupae of H. armigera were induced at high temperatures (33-39 degrees C) with a photoperiod of LD8:16; winter-diapausing and non-diapausing pupae, cultured at 20 degrees C with a photoperiod of LD8:16 and at 27 degrees C, LD16:8, respectively, acted as a control. Retention time of eye spots, weight, and lipid and glycogen levels were compared. At high temperatures, both body weight and energy storage capacity were much higher in summer-diapausing pupae than in non-diapausing pupae reared at 33-39 degrees C. At temperatures (>33 degrees C) high enough to maintain summer diapause, the eye spots of summer-diapausing pupae did not move during the 30-day experiment. However, eye spots of summer-diapausing pupae placed at 30 degrees C began to move about 10 days after they were transferred, significantly later than in non-diapausing pupae reared at 33-39 degrees C or non-diapausing pupae reared at 27 degrees C, which initiated eye spot movement 2 days after pupation. The differences in retention time of eye spots between summer- and winter-diapausing pupae shows that winter diapause is more intense than summer diapause in this insect. The weight loss, and lipid and glycogen metabolism curves indicate that the summer-diapausing pupae's metabolism is very low. We conclude that summer diapause in the cotton bollworm is a true diapause and that the summer diapause enables the cotton bollworm to withstand the high temperatures of summer.