Role of chemical cues from cotton in mediating host selection and oviposition behaviour in Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

We investigated the role of chemoreception in the host selection and oviposition behaviour of Helicoverpa armigera in the laboratory using five cotton genotypes and synthetic volatile terpenes. Female moths oviposited on substrates treated with methanol, ethanol, acetone and pentane extracts of leaves, squares and flowers of the cotton genotypes. Phytochemicals soluble in pentane were the most efficient in eliciting oviposition behaviour. In a two-way bioassay, pentane extracts of leaves or squares of a Multiple Host-plant Resistance genotype (MHR11), Deltapine commercial (DP90), and Smith Red Leaf (SRL) received significantly more eggs than solvent-treated controls. Extracts of squares of the native genotype Gossypium nelsonii did not receive more eggs. Females preferred DP90 and MHR11 to SRL and G. nelsonii. Female moths also laid more eggs on pentane extracts of MHR11 flowers than MHR11 leaves from preflowering, early flowering and peak-flowering plants. In a flight chamber, female moths used olfactory cues at short range to mediate oviposition and discrimination between host plants. Egg-laying, mated females were attracted at a distance (1.5 m) to volatile compounds released by whole plants and odours emanating from filter papers treated with synthetic volatile terpenes. Individually, the terpenes did not stimulate any significant oviposition response. However, there was a significant oviposition response to a mixture of equal volumes of the terpenes ( trans -β-caryophyllene, α-pinene, β-pinene, myrcene, β-bisabolol, and α-humulene). Conversely, antennectomised (moths with transected antennae), egg-laying, mated females did not stimulate any significant oviposition response. The significance of these findings in relation to H. armigera host-plant selection are discussed.     

Efficacy of cashew kernel extracts in the control of the maize weevil,Sitophilus zeamais (Coleoptera: Curculionidae)

Six different solvents namely: methanol, ethanol, water, acetone, pet-ether or n-hexane was used to extract the essential oils from cashew kernel. The extracted oils were evaluated on Sitophilus zeamais (Motschulsky 1855) for mortality, oviposition and adult emergence effects. The long-term storage and water absorption capacity of the treated maize grains was also investigated. Results showed that the steam distillate was most lethal on S. zeamais. The least potent extracts on S. zeamais were those extracted with methanol and ethanol. The extracts drastically reduced oviposition in S. zeamais. Maize grains treated with steam distillate extract had the lowest water absorption capacity, while the highest water absorption capacity was obtained in grains treated with ethanol and methanol. Extracts of steam distillate, n-hexane, petroleum ether and acetone completely prevented infestation and subsequence damage of the treated maize grains for a period of three months. In the grains treated with the methanol and ethanol extracts, damage of 28.61% and 6.24%, respectively were obtained. The results obtained reveal that extracts of cashew kernel are effective in controlling S. zeamais and could serve as an alternative to synthetic insecticides.     

Host plant constituents as oviposition stimulants for a generalist herbivore: European corn borer

Oviposition response of the polyphagous European corn borer,Ostrinia nubilalis (Hübner) (Lepidoptera: Pyralidae), to chemical constituents in host plants was investigated in the laboratory using two-choice bioassays. Foliar extracts of corn (Zea mays L.), pepper (Capsicum annuum L.) and potato (Solanum tuberosum L.) were prepared using the solvents pentane, acetone and methanol. In all three host plants, chemicals soluble in pentane stimulated oviposition. In potato, chemicals extractable in acetone also elicited a positive oviposition response. When presented with a choice between pentane extracts of corn and pepper, females preferred corn. No preferences were exhibited between pentane extracts of corn and potato or pepper and potato. Pentane extracts of corn husks, tassels, silk, and corn leaves from plants at early whorl and tassel (pre-pollen shed) stages of development also stimulated oviposition. Similar extracts from plants at 2-leaf and blister (when kernels resemble blisters) stages were not stimulatory. This indicates that plant phenology affects chemically mediated oviposition response in European corn borer. The potential use of plant chemicals for management ofO. nubilalis in the field is suggested.     

挥发性信息化合物对玉米螟赤眼蜂寄主选择行为的影响

在实验室条件下,利用“Y”型嗅觉仪测定了源于寄主亚洲玉米螟Ostrinia furnacalis (Guenée) 鳞片、卵表以及不同生理阶段的雌蛾附腺的不同浓度的提取物对玉米螟赤眼蜂Trichogramma ostriniae Pang et Chen寄主选择行为的影响,并对其引诱作用大小进行了比较。结果表明： 亚洲玉米螟鳞片的正己烷提取物在0.5 mg/mL和 1 mg/mL浓度时对玉米螟赤眼蜂有明显的引诱作用; 寄主卵的正己烷提取物在5块卵/mL和10块卵/mL浓度时对玉米螟赤眼蜂有显著的引诱作用,而在40块卵/mL浓度时对玉米螟赤眼蜂有极显著的驱避作用; 玉米螟赤眼蜂对亚洲玉米螟交配未产卵和产卵后前期的雌蛾附腺提取液有反应,而对处女蛾和产卵后期的附腺提取液没反应。卵表提取物和附腺提取物比鳞片提取液对赤眼蜂引诱作用强,两者对赤眼蜂的引诱作用没有明显差异。     

Egg distribution patterns in the almond seed wasp,Eurytoma amygdali

The egg distribution patterns ofEurytoma amygdali Enderlein (Hymenoptera, Eurytomidae), which oviposits singly in green, developing almonds, were studied in the laboratory and in the field. In the laboratory, individual females were caged with a number of almonds and the eggs deposited in each fruit were counted. In the field, eggs were censused in almonds of different varieties at regular intervals, over four seasons (1988–91). In the susceptible ‘Retsou’ variety, eggs were uniformly distributed among fruits, both in the laboratory and in the field, as long as the mean number of eggs per almond was ≤2.5. When the mean number of eggs per almond was higher the egg distributions were random. This suggests that, up to a certain level of infestation, females were able to assess egg load of fruits and oviposit in the less infested ones. The main factor enabling the females to discriminate and select the less infested almonds for oviposition is probably a host-marking pheromone. In 1990, the same patterns of egg distribution were observed in samples taken from 5 other almond varieties that are not as susceptible as Retsou and have thicker pericarp and harder endocarp. In 1991 however, when fruits developed more rapidly than in 1990, egg distributions in 3 of these 5 varieties (Truoito, Ai, and Marcona) were not uniform. Although the mean number of eggs per fruit was low (1.2–2.0), many fruits of these varieties contained no eggs. This suggests that, in some less susceptible varieties, egg distribution might also be affected by certain fruit parameters, unfavourable for oviposition, related to the pericarp thickness and endocarp hardness.     

Efficacy of extracts of Melia azedarach L. callus, leaves and fruits against adults of the sweetpotato whitefly Bemisia tabaci (Hom., Aleyrodidae)

Bioassays with extracts of callus, fruit and leaves of the Chinaberry tree, Melia azedarach L., were performed against adults of Bemisia tabaci (Gennadius). Tissue culture of M. azedarach was carried out in Murashige and Skoog (MS) medium supplemented with indole butyric acid and benzyl adenine. Fresh and frozen leaves or fruits and the callus product were extracted in water or methanol at a ratio of 1 : 5 (w/v) for 48 h. The aqueous and methanol extracts were applied on tomato plants to determine their effect on B. tabaci adults and their corresponding oviposition and consequent adult emergence in comparison with the control, distilled water. Results have shown that extracts of callus and different age classes of M. azedarach leaves and fruits have shown significant repellent activity of 58.9–67.7% and have significantly decreased the oviposition rate of the insect without affecting the adult whitefly emergence in comparison with the control. Aqueous and methanol extracts have shown comparable effects against adult whiteflies indicating a similar efficacy of the two solvents in extracting the bioactive components of Melia . Extracts of frozen samples were found to be as effective against the pest as fresh samples, thus allowing storage of Melia parts to be used in case of shortage. This is the first report of the use of M. azedarach callus against B. tabaci; the M. azedarach callus, leaf and fruit extracts seem to present a potential source for management of B. tabaci .     

Evaluation of antimicrobial activity of the lichens <Emphasis Type="Italic">Lasallia pustulata,Parmelia sulcata,Umbilicaria crustulosa</Emphasis>, and <Emphasis Type="Italic">Umbilicaria cylindrica</Emphasis>

The antimicrobial properties of acetone, methanol, and aqueous extracts of the lichens Lasallia pustulata, Parmelia sulcata, Umbilicaria crustulosa, and Umbilicaria cylindrica were studied comparatively in vitro. Antimicrobial activities of the extracts of different lichens were estimated by the disk diffusion test for Gram-positive bacteria, Gram-negative bacteria, and fungal organisms, as well as by determining the MIC (minimal inhibitory concentration). The obtained results showed that the acetone and methanol extracts of Lasallia pustulata, Parmelia sulcata, and Umbilicaria crustulosa manifest antibacterial activity against the majority of species of bacteria tested, in addition to selective antifungal activity. The MIC of lichen extracts was lowest (0.78 mg/ml) for the acetone extract of Lasallia pustulata against Bacillus mycoides. Aqueous extracts of all of the tested lichens were inactive. Extracts of the lichen Umbilicaria cylindrica manifested the weakest activity, inhibiting only three of the tested organisms.     

Semiochemicals from ovaries of gravid females attract ovipositing female houseflies,Musca domestica

Chemical signals originating from the ovaries of gravid females of Musca domestica (Diptera: Cyclorrhapha: Muscidae) attract ovipositing females to common egg-laying sites. Behavioral experiments indicated that females preferred to oviposit in fermented wheat bran containing ovaries from reproductively mature houseflies. Females preferred to oviposit in fermented wheat bran than wet wheat bran. This effect was additive with the attraction to housefly ovaries. Solvent extracts from housefly ovaries were attractive to gravid females. Extracts obtained with hexane were most attractive to gravid females for egg laying, and extracts obtained with ethyl acetate attracted more egg laying than extracts obtained by dichloromethane. Gas chromatography (GC) and gas chromatography coupled with mass spectrometry (GC-MS) analysis showed that tricosane and (Z)-9-tricosene were the main components of the hexane extracts. Both tricosane and (Z)-9-tricosene were shown to elicit dose-dependent aggregation of gravid females in oviposition bioassays, but high doses of either chemical were not attractive.     

家蝇卵巢中的信息化学物质引诱雌蝇产卵的研究

产卵期家蝇MuscadomesticaL .卵巢中的信息化学物质诱引家蝇产卵。行为试验表明 ,雌蝇产卵时优先选择被成熟卵巢污染过的发酵麦麸 ,诱集的产卵百分率依次为 :发酵麦麸 + 1对卵巢 >湿麦麸+ 1对卵巢 >发酵麦麸 >湿麦麸 ;卵巢提取物同样对产卵雌蝇有诱集活力。产卵雌蝇卵巢正己烷、二氯甲烷以及乙酸乙酯的提取物均能诱集雌蝇产卵 ,诱集的产卵百分率依次为 :雌蝇卵巢正己烷提取物 >乙酸乙酯提取物 >二氯甲烷提取物。推测家蝇卵巢中的信息化学物质可能含有家蝇产卵聚集信息素的组分。     

The attractiveness of different odour sources from the fruit–host complex on Leptopilina boulardi, a larval parasitoid of frugivorous Drosophila spp.

In parasitoid insects, successful offspring development depends on the female’s ability to find a suitable host. Specific recognition is often based on responses to olfactory cues, but their source and nature have rarely been determined. –This paper deals with the recognition of odours involved in host location by Leptopilina boulardi[Barbotin, Carton & Kelner-Pillault] (Hymenoptera: Eucoilidae), a larval parasitoid of Drosophila species that develops in mature fruits. The nature and origin of volatile stimuli recognized among odours of the host–fruit complex, and the effect of learning on this recognition, were investigated. Oriented responses to these odours were observed in a four-armed olfactometer and were analysed with the observer software (Noldus Information Technology). Fruit odours alone (banana and pear) were not spontaneously attractive to naive parasitoids, whereas naturally-infested bananas were highly attractive. The attraction was related to the odour that adult Drosophila left on the substrate but not to Drosophila oviposition activity or larval development. A synergism between some fruit odours (banana and pear) and the odour left by adult Drosophila on damp filter paper was observed. However, when testing a non-fruit substrate (mushroom), no synergism was observed. Thus, female L. boulardi may innately recognize host–food substrate odours associated with odours from the adult stage of their host. In addition, an oviposition experience on an infested banana allows L. boulardi females to memorise the fruit odour itself through associative learning. The adaptive significance of this process is discussed.     

Stimuli affecting selection of oviposition sites by female peach twig borer, Anarsia lineatella Zeller (Lepidoptera: Gelechiidae)

In laboratory and field experiments, stimuli were tested that might affect oviposition decisions by female peach twig borer moths, Anarsia lineatella Zeller (Lepidoptera: Gelechiidae). When given a choice between immature green peach fruits, green mature peach fruits and soft-ripe peach fruits, the latter received the fewest eggs. Fuzzy halves of peach fruits received ten times more eggs then shaved hairless halves. Volatiles from both almond and peach shoots induced more oviposition by females than by control stimuli. Similarly, volatiles from immature green peach fruits, mature green or mature hard-ripe peach fruits induced more oviposition than their respective control stimuli. In a choice experiment, volatiles from immature peach fruit stimulated three times more oviposition than those from soft-ripe peach fruit. Discrimination against mature soft-ripe peach fruits as potential oviposition sites may lie in the phenology of A. lineatella and host peach fruits. Larval development to the pupal stage takes 15–27 days. Therefore, any eggs laid on a ripe fruit 14 days before it falls from the tree will not likely develop into adult insects because developing larvae will only reach third or fourth instar before the fruit is decomposed, and only first and second instar larvae can overwinter.     

Olfactometer Responses of Plum Curculio <Emphasis Type="Italic">Conotrachelus nenuphar</Emphasis> (Herbst) (Coleoptera: Curculionidae) to Host Plant Volatiles,Synthetic Grandisoic Acid,and Live Conspecifics

The plum curculio Conotrachelus nenuphar (Herbst) (Coleoptera: Curculionidae) is a major pest of pome and stone fruit, but will also attack other fruits. Males produce the aggregation pheromone grandisoic acid; emitting only the (+)-enantiomer which is attractive to both sexes of the univoltine and multivoltine strains, while the synthetic racemic mixture contains optical isomers with equal amounts of (+)- and (?)-enantiomers. Synergy between odours can increase trap captures and improve monitoring techniques, therefore tests were performed in a dual-choice olfactometer with odours attractive to plum curculios according to literature to determine 1) under what physiological conditions (mating status, age, starvation period) these odours are attractive or repulsive, 2) if the (+)-enantiomer or the odour of live males synergizes with host plant volatiles, and 3) if there is a difference in response between plum curculio strains. Females were exposed to: benzaldehyde; trans-2-hexenal; apples; extracts of: plums, apples, blueberries; grandisoic acid; and live males. Plum essence was found to be the most attractive host-plant odour for both immature and mature virgin females, and immature whole apples were attractive to starved females, while trans-2-hexenal, McIntosh apple essence, benzaldehyde along with the combination of benzaldehyde and plume essence was found to be repulsive. Starvation, age, and mated status all influence response to odours. No synergistic or additive affects were observed between any of the odour combinations tested, including the combination of both the natural and synthetic pheromone and plum essence or apples.     

Divergent responses of two cereal aphids to previous infestation of their host plant

The role of pollen odour in resource location by the pollen beetle, Meligethes aeneus (Fabricius) (Coleoptera: Nitidulidae), a pollen-feeding insect regarded as a pest of oilseed rape, Brassica napus L., (Brassicaceae) crops, was investigated in a linear track olfactometer. Both male and female beetles were attracted to the odour of whole oilseed rape flowers, indicating that these insects can locate their host plants using floral odours as cues. The attractive odour of flowers was found to emanate from all floral parts tested: the petals/sepals, the anthers, and from pollen itself. Therefore, at least part of the attractive odour of oilseed rape flowers emanates from pollen. Beetles were more attracted to floral samples containing anthers than those without anthers when these odours were directly compared in a choice-test, and this indicates that there were detectable differences between them. Anthers and pollen may therefore release distinctive odours that are quantitatively and/or qualitatively different from the odour of the rest of the flower. These experiments support the hypothesis that pollen-seeking insects use pollen odour cues to locate this food source.     

Response of female Lutzomyia longipalpis to host odour kairomones from human skin

Lutzomyia longipalpis Lutz & Neiva (Diptera: Psychodidae) is the vector of Leishmania chagasi, the aetiologic agent of visceral leishmaniasis in the New World. In the present study, the response of female sandflies from Jacobina, Brazil, to human odours from six different volunteers was investigated. Glass Petri dishes were handled by different volunteers and then exposed to female sandflies. There was a significant difference between subjects in that some individuals were more attractive or less repellent to sandflies. Response of flies to handled Petri dishes was higher during the first minutes of observation, suggesting the presence of volatile compounds in hand odours. Extracts of glass Petri dishes that had been handled by the volunteers were made with organic solvents such as acetone, methanol, pentane and ether. These were then concentrated and tested for sandfly response. Only extracts carried out with non-polar solvents such as pentane and ether were able to transfer odours from handled glass Petri dishes onto clean dishes. The attractivity of male and female human subjects was monitored for 80 days, and minor fluctuations in attractiveness were observed.     

Increasing the availability of food and the reproduction ofPoecilus cupreus in a cereal field by strip-management

The spruce bud moth,Zeiraphera canadensis Mutt & Free. (Lepidoptera: Tortricidae), displayed a bimodal activity pattern in two young white spruce,Picea glauca (Moench) Voss, plantations. During the day, when temperatures were highest and relative humidities lowest, most moths remained in the lower crown, presumably in response to the risk of hygrothermal stress and/or predation. Flight activity peaks occurred near dawn and dusk. Moths moved higher in the crowns shortly after sunset and many were observed flying above the tree canopy. This activity peak was associated with oviposition and mating. Oviposition started at sunset and continued for approximately 3 h. Females probed oviposition sites with their ovipositor before ovipositing, suggesting that they use sensillae on the ovipositor to evaluate potential oviposition sites. Most mating occurred between 22∶00 and 04∶00 h. Males hovered adjacent to branches containing females before alighting. After alighting, males usually approached to within 2–5 cm of females and then remained motionless for a short time before attempting mounting, indicating that some short-range communication usually occurs prior to mating. Reduced flight activity between midnight and dawn was not due to low temperatures, which were above the minimal temperatures for flight for males (10°C) and females (12°C) determined in the laboratory.     

Response of cowpea bruchid,Callosobruchus maculatus (Fab.) (Coleoptera: Chrysomelidae) to cheese wood,Alstonia boonei De Wild stem bark oil extracted with different solvents

Cheese wood, Alstonia boonei De Wild stem bark was extracted with five different solvents namely: methanol, ethanol, acetone, petroleum ether and n-hexane. The extracts were tested for insecticidal activity on cowpea bruchid, Callosobruchus maculatus (Fab.). Adult mortality and adult emergence of the insects were investigated. Oviposition deterrences using dual-choice and multiple-choice tests were also investigated as well as percentage damage and weight loss at temperature of 28?±?2?°C and 75?±?5% relative humidity. Results showed that at the rate of 2% extract per 20?g of cowpea seeds, methanol, petroleum ether and n-hexane extracts caused 100% mortality of adult cowpea bruchid after four?days of post treatment. When the bruchid had choice of oviposition substrate, they laid significantly (p?<?0.05) fewer eggs on cowpea seeds treated with extracts compared to untreated seeds. There was no adult emergence in seeds treated with methanol, ethanol, petroleum ether and n-hexane extracts and 100% reduction in F₁ progeny was recorded compared with untreated that had 81.86% adults emergence and 0% reduction in F₁ progeny. The results obtained from this study revealed that methanol, ethanol, acetone, petroleum ether and n-hexane extracts of A. boonei stem bark were effective in controlling C. maculatus and could serve as an alternative to synthetic insecticides for the protection of stored cowpeas against bruchids.     

Behavioural and electrophysiological responses of grape berry moth (Lep., Tortricidae) to selected plant extracts

Abstract:  Four non-host plant extracts from Bifora radians , Arctium lappa , Humulus lupulus and Xanthium strumarium , were tested against adult grape berry moths, Paralobesia viteana , for their potential as repellents or oviposition deterrents. Responses were compared with those elicited by the major component of the P. viteana sex pheromone. Moths of both sexes exhibited varying electroantennogram (EAG) responses from 1.7 to 2.4 mV to volatile odours from plant extracts, with the greatest response to extracts of H. lupulus (2.4 mV). A multi-arm olfactometer was used to assay the behavioural response of moths to the same treatments. Male moths responded significantly to H. lupulus extract, although their strongest response was to the sex pheromone (30.0%). Female moths showed no behavioural response to the sex pheromone, attracting only 6.7% of moths, but they were attracted to extracts of H. lupulus , X. strumarium and A. lappa , with 25.0%, 21.7% and 15.0% of the released insects responding to these treatments, respectively. Choice tests were conducted to compare oviposition by P. viteana on untreated grapes and on grapes treated with one of the extracts. Despite the attraction in the olfactometer assays, H. lupulus extract significantly reduced egg laying; only 12.5% of the total deposited eggs were laid on berries treated with this extract. The extract of B. radians provided complete protection against oviposition.     

Mise en évidence d'une activité kairomonale des œufs de la pyrale du maïs sur les femelles deTrichogramma maidis

Résumé Des extraits de la surface d'œufs de Pyrale du ma?s ont été testés en microenceinte et dans un olfactomètre linéaire sur des ♀♀ deTrichogramma maidis. En microenceinte, les extraits stimulent la locomotion et le comportement exploratoire des ♀♀. En olfactomètre ils stimulent à distance la remontée des tubes vers la source. Ces résultats impliquent la présence de substances à activité kairomonale sur les œufs de Pyrale du ma?s. Des analyses préliminaires des extraits ont été faites par chromatographie en phase gazeuse couplée ou non à la spectrométrie de masse et démontrent la présence de nombreux hydrocarbures.

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Summary Extracts from the surface of eggs of the European corn borer were tested in small observation chambers and in a linear olfactometer for their effects on the searching behaviour of the egg-parasitoidTrichogramma maidis. In the observation chambers, the extracts elicited exploratory behaviour by female parasitoids with no previous oviposition experiences and increased the time the females spent in walking and drumming their antennae. Recording the position of 10 ♀♀ each minute for 15 min in 2 tubes of an olfactometer, indicated that upwind motion was significantly greater in the tube containing 10 μl of an egg extract than in the control tube containing just solvent. It is concluded that kairomones are present onO. nubilalis eggs and could act as searching incitants and short-range attractants. Preliminary GC and GC/MS analysis revealed the abundance of hydrocarbons in the extracts but the active chemicals were not characterized.

     

In vitro screening of seaweed extract on the proliferation of mouse spleen and thymus cell

A total number of 31 types of seaweed were assessed with regard to their effects on the proliferation of mouse spleen and thymus cells in a culture, using an MTT reduction assay. Acetone:dichloromethane (1∶1) extracts of three seaweed plants:Derbesia marina, Sargassum sp., andHisikia fuziformis, exhibited significantly positive effects on the survival of mouse spleen and thymus cellsin vitro. The acetone: dichloromethane (1∶1) extracts ofSargassum sp., in particular, much more potent effects on thymus cell activation than did any of the other types of seaweed. However, the methanol extracts ofSargassum ringgoldianium andChondrus crispus exerted a stimulatory influence only on the proliferation of mouse spleen cells, whereas the methanol extracts ofGrateloupia lanceolata exhibited significant cell proliferation properties in both spleen and thymus cells.     

Mosquito larvicidal activity of aqueous extracts of long pepper (Piper retrofractum vahl) from Thailand.

Aqueous extracts of nine medicinal plants were bioassayed against larvae of Culex quinquefasciatus Say and Aedes aegypt (L.). Among these plants, the long pepper, Piper retrofractum Vahl (Piperaceae), showed the highest level of activity against mosquito larvae. To gain more information on larvicidal activity of P. retrofractum, fresh fruits of this plant were extracted in water and the extracts made into powder and bioassayed against 3rd and 4th instar larvae of Cx. quinquefasciatus and Ae. aegypti in the laboratory. Extracts of unripe (001/3) and ripe (002/3 and 001/4) fruits showed different levels of activity against Cx. quinquefasciatus larvae. Extracts 001/3 and 002/3 were equi-toxic to a Bacillus sphaericus resistant and susceptible strains, both from Thailand. The ripe fruit extract 002/3 was somewhat more active against Ae. aegypti than Cx. quinquefasciatus. Another ripe fruit extract (001/4) was much more toxic to both mosquito species. Diluted solutions of the solid extract (002/3) in distilled water lost their larvicidal activity upon aging. Loss of activity at 25 degrees C was greater than that stored at 4 degrees C, and greater in water than in acetone solution.