Effect of Bacillus thuringiensis naturally colonising Brassica campestris var. chinensis leaves on neonate larvae of Pieris brassicae

The feeding of neonate larvae of Pieris brassicae (Order Lepidoptera) on leaves of brassica plants that had been colonised by Bacillus thuringiensis resulted in the death of 35% of the population within 72 h. The bacteria multiplied in the cadavers, resulting in an increase of about 50-fold compared to the living insects. Surviving insects showed no ill effects during the time of the study. There was negligible multiplication of B. thuringiensis in the frass.     

Genetische Untersuchungen mit Gartenkohl (Brassica oleracea)

Ohne ZusammenfassungMit Tafel 3–10 und 13 Textfiguren     

The effect of ozone fumigation and different Brassica rapa lines on the feeding behaviour of Pieris brassicae larvae

Elevated levels of tropospheric ozone and their effects on plants have been studied for a great number of years. Ozone is a gaseous pollutant and acts as a phytotoxin. Even though ozone is known to change the physiology of plants, little attention has been given to the indirect effects of ozone on plant-insect interactions. This paper addresses this question by investigating the interactive effects of ozone and plant genotype on insects. Lines of rapid-cycling Brassica rapa (L.) selected for their contrasting sensitivity to ozone and Pieris brassicae (L.) (Lepidoptera: Pieridae) were used as a model system. The effect of differences in ozone sensitivity and ozone fumigation on the plant's carbon and nitrogen pools, the feeding preference, and behaviour of P. brassicae larvae were investigated. The results show that the plant's susceptibility to ozone interacts in a complex way with ozone induced alterations in the suitability of the plant for the insect. Only the larval performance on the sensitive line was affected by ozone exposure. Biochemical changes in the resistant B. rapa line made the plant a better food source for the insects, since the digestibility of this plant was significantly higher than that of the sensitive line, and the larvae pupated more quickly and were heavier.     

Diuresis in the cabbage white butterfly, Pieris brassicae: Fluid secretion by the malpighian tubules

The sudden weight loss at the pupal-adult ecdysis of the cabbage white butterfly, Peiris brassicae, is caused by a diuresis which greatly reduces the blood volume after emergence. Study of the Malpighian tubules as isolated preparations shows that only those of the adult insect possess the ability to secrete fluid at extremely fast rates when stimulated. In contrast to the fast-secreting tubules of bloodsucking insects, the mechanism of fluid secretion is not unusual, probably depending on the active transport of potassium ions.     

The influence of temperature on respiration of diapausing pupae of Pieris brassicae (Lepidoptera)

The respiration of diapausing Pieris pupae has been measured at different temperatures between 5 and 35°C in animals maintained at 20°C, either 14 or 74 days after larvo-pupal ecdysis or at 5°C for 30 or 60 days.

The sudden transfer of animals from 5 to 15, 20, 25, 30, 35°C or from 20 to 30, 35°C results in a respiratory overshoot whose characteristics (duration, height, extra-respiration) depend on experimental conditions.

After a certain period of acclimation, overshoots are eliminated. The respiratory rate except for animals maintained during 74 days at 20°C can then be represented as an exponential function of temperature.

The Q₁₀ values change according to the treatment given to pupae.

The respiratory rate of male pupae is higher than that of female ones.

The following points are discussed:

1. 1.|The meaning of overshoots is analysed according to economy and metabolic homeostasis, showing the existence of acclimation.

2. 2.|Exponential curves which are not relevant to non-diapausing pupae or to the diapausing ones taken at larvo-pupal molting are characteristic of steady metabolism. These curves can be interpreted as the result of the temperature effect on a master respiratory reaction which would then be rate limiting.

3. 3.|Wintering leads to gradual and slow adaptation to cold temperatures which brings both a respiratory increase, a decrease of the Q₁₀ and of the activation energy of the master reaction.

青花菜谷胱甘肽-S-转移酶基因克隆及其表达分析

为探讨青花菜在模拟酸雨胁迫下谷胱甘肽-S-转移酶的表达变化,克隆了青花菜谷胱甘肽-S-转移酶基因(glutathione-S-transferase,GST)的cDNA序列全长,并进行了生物信息学和表达分析。结果表明:青花菜GST基因cDNA全长为915bp,开放阅读框为642bp,编码213个氨基酸,推测分子式为C1091H1719N289O306S5,分子量为23 940.7,没有跨膜螺旋区域和信号肽。系统进化树分析表明,该青花菜基因GST与芥菜的GST聚类关系最近。实时荧光定量PCR结果显示,在模拟酸雨胁迫下,GST基因的表达量在胁迫初期显著增大,随时间延长开始下降,表明其参与了青花菜抗酸雨的应答反应。     

Developmental instability in cabbage aphid (Brevicoryne brassicae) populations exposed to heavy metal accumulated host plants

Random deviations from the perfect symmetry of normally bilaterally symmetrical characters exist during individual development as a result of various environmental conditions. Fluctuating asymmetry (FA) is often used as a measurement of developmental instability, and within-environmental variation (CV_e) is also considered as an indicator of developmental deviations. These two parameters may indicate the quality of the environmental habitat of organisms. For herbivore insects, such as aphids, any change in their host plants conditions is important and directly affects their development. The presented investigation revealed that both Lead (Pb) and Copper (Cu) accumulation in different host plants resulted in a significant amount of deviations from bilateral symmetry in cabbage aphid (Brevicoryne brassicae). Cabbage aphid populations showed higher FA and CV_e on heavy metal accumulated cabbage and radish than on non-contaminated cabbage and radish plants. However, the pattern between developmental instability and fitness measurements was inconsistent. Thus, this study does not support the hypothesis that asymmetry is a valuable bioindicator of developmental instability.     

Effects of different host plants and rearing atmosphere on life cycle of large white cabbage butterfly,Pieris brassicae (Linnaeus)

We studied the effect of different host plants and rearing atmosphere on life cycle of cabbage butterfly, Pieris brassicae. Insects were reared in the field (fluctuating weather) as well as in the laboratory (constant rearing conditions) on four host plants, viz. cabbage, cauliflower, knol-khol and broccoli. Significant differences were not found in the incubation and pupal period of butterfly reared on different host plants. However, larval period was found to be significantly lower on cabbage followed by knol-khol and highest on broccoli. Therefore, the developmental period (from eggs to adult) was reasonably lower on cabbage. Furthermore, inverse relationship was found in the body weight of butterfly and developmental period, where weight of full grown caterpillar and pupae was significantly higher on cauliflower and lowest on cabbage. Besides, significant differences were not found in the body weight of P. brassicae caterpillar reared under field and in the laboratory. Nevertheless, pupal weight of butterfly was significantly higher under field conditions than the laboratory conditions. Overall, the development of P. brassicae was much faster on cabbage than other hosts; but its body weight was considerably higher on cauliflower.     

Stimulation of oviposition by the cabbage root fly Erioischia brassicae

A study was made of physical and chemical aspects of oviposition stimulation in E. brassicae.It was shown that contact chemostimulation provided by host plant material stimulated the gravid female to probe with the ovipositor into cracks in which eggs were laid. Carbon disulphide stimulated oviposition at certain concentrations in water, but solutions of carbon disulphide and three other chemicals previously shown to stimulate oviposition were less effective in mixtures than the juice of swede, Brassica napus var. napobrassica, a favoured host. The addition of mangold juice or 20% sucrose solution to solutions of chemostimulants appeared to enhance oviposition.The most favourable soil for oviposition was of particles approximately 1 mm in diameter which may just permit the complete entry of the extended ovipositor, and provide irregularities against which eggs can be laid.

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Zusammenfassung Eine Untersuchung der physikalischen und chemischen Aspekte der Eiablage-Stimulation von Erioischia brassicae wurde durchgeführt.Es wurde gezeigt, daß chemischer Kontakt mit dem material der Wirtspflanze die trächtigen Tiere dazu veranlasste, Vertieferungen mit dem Ovipositor abzutasten, in welche dann Eier abgelegt wurden.Gewisse Konzentrationen wässriger Schwefelkohlenstofflösungen stellten einen Anreiz zur Eiablage dar. Jedoch waren Mischungen von CS₂-Lösungen mit vier anderen Reizstoffen weniger wirkungsvoll als der Saft des bevorzugten Wirtes, Brassica napus L. var napobrassica (L.) Rchb. (Steckrübe). Der Zusatz von Mangoldsaft oder 20%-iger Zuckerlösung zu Lösungen von chemischen Reizstoffen schien die Eiablage zu steigern.Der bevorzugte Boden für die Eiablage hatte eine Korngrösse von ungefähr 1 mm Durchmesser, welche wahrscheinlich das vollständige Eindringen des ausgestreckten Ovipositors gestattet und das günstigste Substrat zur Eiablage darstellt.

     

结球甘蓝下胚轴原生质体培养再生植株体系的优化研究

以结球甘蓝‘新夏50’的无菌苗下胚轴为材料,对影响原生质体分离、纯化与培养的主要因素进行研究,建立适合结球甘蓝原生质体游离、纯化、收集、培养以至再生出完整植株的实用技术体系,为其非对称细胞融合及品种改良与创新等研究奠定基础。结果表明:2.5%纤维素酶R-10+0.05%果胶酶Y-23+9CPW+5mmol/L MES的混合酶液,从4d苗龄的下胚轴上分离出高产率的原生质体。在改良B5+0.5mg/L 2,4-D+0.2mg/L 6-BA+0.2mg/L NAA的液体培养基上,原生质体分裂旺盛。形成愈伤组织后经芽诱导和生根培养,获得了再生植株。倍性检测结果表明,不同原生质体所获得的24株再生植株中,19株为正常二倍体,4株为嵌合体,1株为四倍体。     

Effect of host plant odour on the behaviour of the adult cabbage root fly, Erioischia brassicae

Laboratory experiments showed that the odour of leaves of host plants stimulated an increase in activity in gravied female E. brassicae while the odour of non-host foliage had little effect. Males and non-gravid females showed very little response to the odour of both types of foliage. Other experiments failed to show any orientation of the flies towards sources of host plant odour, although more flies were caught in traps containing the juice expressed from a host plant than in those containing water. It is suggested that an unoriented activation of gravid female E. brassicae by host plant odour enhances their chance of being caught in traps emitting the odour, and of making contact with host plants in the field.

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Zusammenfassung Laborversuche zeigten, daß der Blattduft der Wirtspflanzen die Aktivität der trächtigen Weibchen von Erioischia brassicae steigert, während der Duft von Fremdlaub nur einen geringen Effekt hervorrief. Die Reaktion der männlichen und der jungen weiblichen Tiere auf den Duft beider Laubtypen war gleichmäßig schwach. Weitere Laborversuche zeigten keine Orientierung der Tiere auf die Quelle des Wirtspflanzenduftes, dennoch wurden mehr Tiere in Fallen gefangen, welche Wirtspflanzensaft enthielten als in solchen, die nur mit Wasser beschickt waren. Es wird angenommen, daß die trächtigen Weibchen von Erioischia brassicae durch den Wirtspflanzenduft unorientiert aktiviert werden und somit für sie eine grössere Möglichkeit besteht, in Fallen gefangen zu werden, die einen solchen Duft ausströmen, und in der Natur auch mit Wirtspflanzen in Kontakt zu kommen.

     

Perception of the oviposition deterring pheromone by tarsal and abdominal contact chemoreceptors in Pieris brassicae

Perception of the oviposition deterring pheromone by contact chemoreceptors in female Pieris brassicae was studied employing a tip recording technique. Electrophysiological responses of tarsal taste hairs to eggwash solutions show a marked increase in frequency of spikes originating from only one sensory cell. This suggests that in foretarsal taste hairs females, apart from the glucosinolate cells also possess sense cells specifically sensitive to the oviposition deterring pheromone.Morphological studies by means of the scanning electron microscope revealed that the ovipositor of P. brassicae is provided with two groups of contact chemoreceptors. Electrophysiological recordings from these sensilla indicate the presence of at least three sensory cells, one of them being a mechanoreceptor. Stimulation with eggwash evokes a slight increase in spike frequency which cannot be ascribed to one particular sense cell. This indicates that abdominal taste hairs in some way may participate in the perception of the oviposition deterring pheromone. Responses to glucosinolates do not differ significantly from control stimulations.

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Résumé Une technique d'enregistrement apical a été utilisée pour examiner la perception d'une phéromone dissuadant la ponte par les poils récepteurs chimiques de contact des femelles de Pieris brassicae. Les réponses électrophysiologiques des poils gustatifs des tarses en présence de solutions de rinçage d'oeufs présentent une fréquence marquée des potentiels d'action provenant principalement d'une cellule sensorielle. Ceci suggère que les poils gustatifs des tarses des pattes antérieures des femelles possèdent, en plus de cellules répondant aux glucosinolates, des cellules sensorielles sensibles spécialement à la phéromone dissuadant la ponte.Des études morphologiques au microscope à balayage révèlent que l'oviposition de P. brassicae est pourvu de deux groupes de chimiorécepteurs de contact. Des enregistrements électrophysiologiques de ces sensilles révèlent la présence d'au moins trois cellules sensorielles, l'une d'entre elles étant un mécanorécepteur. La stimulation avec la solution de rinçage des oeufs évoque un léger accroissement de la fréquence des potentiels d'action qui ne peut être attribué à une cellule sensorielle particulière. Ceci indique que les poils gustatifs abdominaux peuvent participer d'une certaine façon à la perception de la phéromone dissuadant la ponte. Les réponses aux glucosinolates ne diffèrent pas significativement des stimulations témoin.

     

Visual learning in assays of sinigrin solution as an oviposition releaser for the cabbage butterfly, Pieris rapae

Pieris rapae (L.) oviposited readily on paper wetted with sinigrin solution, if it was yellow, green, light blue or white, but not if violet, red, or black. On violet discs, feeding responses distracted the butterflies from oviposition and they often laid on neighbouring green discs which lacked sinigrin. When some discs with water and others with sinigrin solution were offered together, butterflies oviposited more on water discs of the same colour as the sinigrin discs, as opposed to those of a different colour. Within a range of sinigrin solutions tested (10–1000 ppm) the higher concentrations elicited most oviposition. Preference was slight for different concentrations applied to physically identical discs, but marked for the higher of two concentrations (100 vs 75 ppm) on discs of different but equally acceptable appearance. It was concluded that by trial-and-error, ovipositing butterflies learnt to associate the appearance of the oviposition site with its contact chemical stimuli, and that learning either increased or decreased the specificity of responses to sinigrin-treated discs, according to the design of the assay. Learning influenced preference differently in butterflies offered two as opposed to three kinds of acceptable site.

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Résumé La ponte de Pieris rapae L., phytophage spécialiste de crucifères, a été étudiée avec des rondelles de papier buvard coloré, imbibées avec une solution de sinigrine, substance spécifique, non volatile, des plantes hôtes. Les papillons ont bien pondu lors de répétitions sur papiers jaune, vert, bleu clair ou blanc, et peu sur papiers rouge, violet ou noir. Avec des rondelles violettes, la prise d'aliments a distrait de la ponte, et des oeufs ont alors été souvent pondus sur des rondelles vertes proches sans sinigrine. Quand des rondelles avec de l'eau et d'autres avec des solutions de sinigrine ont été offertes simultanément, les papillons ont pondu plus sur les rondelles imbibées d'eau de la même couleur que celles avec de la sinigrine, que sur celles d'une autre couleur. Dans la gamme des concentrations de solution de sinigrine examinées (10 à 1000 ppm les plus élevées ont induit les pontes les plus importantes. Il y a eu peu de préférence entre des rondelles de même couleur ayant des concentrations différentes, mais le choix a été net quand les rondelles étaient de couleurs différentes et pourtant acceptables, dans ces conditions la concentration la plus élevée (100 versus 75 ppm) était préférée. On peut en conclure que les papillons, par un apprentissage avec essai et erreur, lors de la ponte ont associé l'aspect du lieu de ponte avec les stimulations chimiques de contact; l'apprentissage a, soit augmenté, soit réduit la spécificité des réponses aux rondelles traitées à la sinigrine, suivant les conditions de l'expérience. La discussion examine la signification du tambourinage par les pattes antérieures comme séquence préliminaires de la ponte.

     

The effect of sinigrin on the feeding of Pieris brassicae L. larvae transferred from various diets

It is well known that mustard oil glucosides act as feeding stimulants for P. brassicae larvae. However it is shown here that the larvae will feed on a diet which contains no mustard oil glucoside if they are placed on it from the time of hatching. Such larvae complete their development a little more slowly than those on a diet containing a glucoside. When transferred to a diet which contains powdered cabbage or sinigrin their feeding is increased by about 20 per cent. Larvae which have been reared on fresh cabbage will not accept a diet even if it contains sinigrin but larvae which have been reared on diets containing dried cabbage or sinigrin feed much better on either of these diets than on one containing neither.

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Zusammenfassung Es ist wohl bekannt, daß Senfölglukoside als Fraßstimulatien für Raupen von Pieris brassicae L. wirken. Hier wird jedoch gezeigt, daß die Raupen von einer Nahrung fressen, die keine Senfölglukoside enthält, wenn sie vom Schlüpfen an auf ihr gehalten werden. Solche Larven vollenden ihre Entwicklung ein wenig langsamer als die, welche an glukosidhaltiger Nahrung aufgezogen werden. Wenn sie auf eine Nahrung übertragen werden, die gepulverten Kohl oder Sinigrin enthält, wird ihre Nahrungsaufnahme um etwa 20% gesteigert. Raupen, die an frischem Kohl aufgezogen wurden, nehmen künstliche Diät auch dann nicht an, wenn sie Sinigrin enthält. Aber Raupen, die mit Nahrung aufgezogen wurden, welche getrockneten Kohl oder Sinigrin enthält, fressen sehr viel besser an einer dieser Diäten als an einer, die keines von beiden enthält.

     

Life table parameters of large white butterfly Pieris brassicae (Lepidoptera: Pieridae) on different cabbage varieties

Effect of four different cole crops (Brassica oleracea var. botrytis, Brassica oleracea var. capitata, Brassica oleracea var. italica and Brassica oleracea var. viridis) on biological parameters of the large white butterfly Pieris brassicae was evaluated at temperature 26 ± 1 °C, 60 ± 5% R. H. and a photoperiod of 16: 8 (L:D) h. The shortest larval and pupal period stages were recorded on B. oleracea var. botrytis (22.18 ± 0.20 days) and (13.32 ± 0.17 days), respectively. The life span was longest on B. oleracea var. viridis (60.43 ± 2.34 days) and shortest on B. oleracea var. botrytis (50.19 ± 0.51 days). The highest percentage of larval and pupal mortality was observed on B. oleracea var. viridis (74%), and (53%), respectively. We found that P. brassicae prefers B. oleracea var. botrytis and B. oleracea var. capitata among cole crops; it is due to the lowest percentage of larval and pupal mortality and the highest rate of life table parameters, including survival rate (lx) and life expectancy (ex), which makes them to be susceptible varieties to this pest. Contrary, a longer developmental time on B. oleracea var. viridis may be attributed to poor nutritional status and reduced survival of the cohort, resulting in high rates of mortality, which was partial resistance to pest. Knowledge of the biology and life table parameters of P. brassicae on different cole crops could be effective in detecting and monitoring the pest infestation, variety selection and crop breeding.     

Immunomodulatory activity of Sulforaphane, a naturally occurring isothiocyanate from broccoli (Brassica oleracea)

The effect of Sulforaphane on the immune system was studied using BALB/c mice. Intraperitoneal administration of five doses of Sulforaphane (500 microg/dose/animal/day) was found to enhance the total WBC count (12,950 cells/mm3) on 9th day. Bone marrow cellularity (23 x 10(6) cells/femur) and number of alpha-esterase positive cells (1346.66/4000 cells) were also increased by the administration of Sulforaphane. Treatment with Sulforaphane along with the antigen, sheep red blood cells (SRBC), produced an enhancement in the circulating antibody titre and the number of plaque forming cells (PFC) in the spleen. Maximum number of PFC (315.83 PFC/10(6) spleen cells) was obtained on the 6th day. Administration of Sulforaphane also showed an enhancement in the phagocytic activity of peritoneal macrophages. Moreover administration of Sulforaphane significantly reduced the elevated level of TNF-alpha production by LPS stimulated macrophages. These results indicate the immunomodulatory activity of Sulforaphane.     

Direct somatic embryogenesis from pericycle cells of broccoli (Brassica oleracea L. var. italica) root explants

Cotyledon, hypocotyl or root explants of 7-day-old broccoli seedlings were cultured on Murashige and Skoog (MS) agar or liquid medium supplemented with 1.0 mg l⁻¹ 2,4-dichlorophenoxyacetic acid (2,4-D). The frequency of direct somatic embryo formation was 100% when root explants were cultured in liquid medium. Histological analysis indicated that somatic embryos were initiated directly from the pericycle cell layers of root explants as early as 1 day after liquid culture. Genotype did not affect the frequency of somatic embryo formation or the number of somatic embryos per explant. All broccoli genotypes examined had 100% somatic embryo induction efficiency, and the number of somatic embryos per 0.8 mm root segment ranged from 22.9 in ‘Luhui’ to 26.0 in ‘Haizi’. The number of normally developed somatic embryos in culture increased with increasing 2,4-D concentration. Plantlet regeneration frequency was the highest (73.3%) when germinated plantlets were transferred to 1/2 strength MS agar medium containing 1.0 mg l⁻¹ 6-benzyladenine (BA). When regenerated plantlets were transferred to a greenhouse, approximately 75% survived and there were no morphological differences between regenerated plants and seed-derived controls. The protocols established in this study will benefit large-scale vegetative propagation and transformation-based genetic improvement of broccoli.     

Diuresis in the cabbage white butterfly, Pieris brassicae: Water and ion regulation and the rôle of the hindgut

The significance of diuresis in the water and ion balance of newly emerged Pieris was examined by comparing the composition of haemolymph and urine during diuresis. The high potassium content of the urine results in a marked increase in the Na/K ratio of the haemolymph. The haemolymph osmolarity is well regulated, in spite of the very hypo-osmotic urine. By means of an isolated preparation of the ileum, it was shown that rapid resorption of potassium ions by this part of the hindgut is responsible for the low osmolarity of the urine.     

Observations on adult weight and wing area in Plusia gamma L. and Pieris brassicae L. In relation to larval population density

Crowding of lepidopterous larvae can influence adult morphology and physiology. With P. gamma and P. brassicae increasing emergence weight was not accompanied by proportionately as great increases in wing area. Larval crowding decreases both weight and wing area and accentuates thedisproportion in the relation between wing area and weight. In the two species fore- and hind-wings are differently affected and the sexes were also affected differently. A warmer season also decreases fore- and hind-wing areas but distinct differences exist in the relative extent of the decreases. To obtain adequate data, both fore- and hind-wings must be measured.

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Résumé Le groupement chez les larves de lépidoptères provoque la réduction du poids et des dimensions des ailes chez les adultes, et il est possible qu'elle contribue à provoquer la migration par son effet sur la morphologie et la fonction. Le rapport entre le poids total et la surface des ailes est souvent appelé wing loading et on sait que ce rapport croît avec l'accroissement de poids des différentes espèces animales qui volent.L'effet du groupement larvaire sur le poids du corps et le wing loading chez Plusia gamma L. et Pieris brassicae L. a été étudié en comparant les adultes nouvellement éclos d'élevage de larves solitaires et de larves groupées. Le groupement réduit le poids et la surface des ailes d'une façon non proportionnelle, mais toutefois décroissante, wing loading. Pour chaque espèce et chaque sexe, le wing loading décroît pour un poids à l'éclosion décroissant, et ceci explique partiellement l'effet du groupement sur le wing loading.Chez P. gamma l'aile antériure était plus grande que l'aile postérieure. Toutefois l'aile antérieure était la plus réduite en dimensions par le groupement, l'effet étant le plus marqué chez la femelle. La température affectait aussi le poids et la surface des ailes. Dans les élevages d'individus isolés, des températures plus élevées réduisaient principalement le poids des mâles, et la surface de leurs ailes postérieures, tandis que les surfaces des ailes antérieures et postérieures des femelles étaient plus affectées, de sorte que le wing loading décroissait chez les mâles et augmentait chez les femelles.Chez P. brassicae l'aile postérieure est la plus grande et décroissait davantage avec le groupement. Le groupement ne paraissait pas affecter différemment les deux sexes, mais les résultats semblaient indiquer que l'effet pouvait être accentué chez les mâles et diminué chez les femelles quand les températures baissaient. Comme chez P. gamma des températures plus élevées diminuaient le poids et la surface des ailes chez P. brassicae mais du fait que la surface des ailes était la plus affectée, le wing loading augmentait chez les deux sexes. Le groupement, cependant, accentuait l'effet de la température sur le poids tandis qu'elle réduisait le wing loading. ainsi chez les deux espèces, les réductions du poids et de la surface des ailes dûes à l'élévation de la température ne sont pas identiques avec les réductions dûes au groupement.Les expériences ont montré également que, alors que les mesures directes de certaines dimensions pouvaient présenter une précision adéquate, les renseignements sur les ailes antérieures et postérieures devaient être groupés.