Tyramine from the leaves of wild parsnip: a stimulant and synergist for oviposition by the black swallowtail butterfly

Abstract .Oviposition stimulants from the foliage of wild parsnip, Pastinaca sativa (Apiaceae), were isolated by column chromatography and HPLC and tested in bioassay experiments with hand-held female black swallowtail butterflies, Papilio polyxenes (Lepidoptera: Papilionidae). Two of the stimulants were identified as tyramine and trans -chlorogenic acid. A combination of tyramine, trans -chlorogenic acid and an active neutral fraction was needed to elicit a significant oviposition response. These results are discussed in the context of previous research on the oviposition stimulants of swallowtail butterflies and on the significance of tyramine as a neuromodulator of physiological processes in invertebrates.     

Tarsal contact chemoreceptors of the black swallowtail butterfly Papilio polyxenes: responses to phytochemicals from host- and non-host plants

Abstract Tarsal contact chemoreceptors of the black swallowtail butterfly Papilio polyxenes (Lepidoptera: Papilionidae) were stimulated with leaf-surface extracts and ethanolic extracts of whole leaves of a host-plant (Daucus carota) and a non-host (Brassica oleracea). Both leaf extracts evoked large numbers of spikes but stimulated different receptor neurones. It is concluded that a large difference exists between the sensory responses to host and non-host extracts. Two chemicals, luteolin 7-0-(6"-0-malonyl)-β-D-glucopyranoside and trans- chlorogenic acid, known to be present in the host and known oviposi-tion stimulants for P.polyxenes , were also tested and shown to be active. No responses were found to luteolin 7-O-β-D-glucoside or to luteolin 7-O-β-D-glucuronide. These flavonoids occur in D.carota foliage, but do not stimulate oviposition.     

Effects of plant odor on oviposition by the black swallowtail butterfly,Papilio polyxenes (Lepidoptera: Papilionidae)

Black swallowtail females laid more eggs on plant models treated with contact stimulants and volatiles from carrot leaves than on models treated only with contact stimulants. The volatiles enhanced landing rates and females alighted more frequently on artificial leaves treated with host volatiles than on adjacent control leaves. Volatiles from cabbage, a nonhost, inhibited landing rates on artificial leaves treated with carrot contact stimulants. Examination of antennae revealed two major types of sensilla, believed to be olfactory in function. Electroantennogram preparations responded more strongly to carrot volatiles than to cabbage volatiles and several shared responses at particular retention times to carrot volatile components eluting from a gas chromatograph. Our results are consistent with a long-standing hypothesis that behavioral responses to essential oil components characteristic of the larval food plants have facilitated host shifts in the genus Papilio.     

Food utilization and antioxidant enzyme activities of black swallowtail in response to plant phototoxins

Phototoxic compounds are widely distributed among plant families; due to their ability to bind covalently to DNA or to react with oxygen and generate toxic oxyradicals, these compounds are toxic to a variety of herbivorous organisms. Black swallowtail (Papilio polyxenes) larvae feed exclusively on phototoxic host plants in the Apiaceae and Rutaceae. In this study, we examined the toxicity of four phototoxins—three furanocoumarins and one β-carboline alkaloid—to P. polyxenes, as well as the inducibility of antioxidant enzyme defenses in response to these phototoxins. Neither the furanocoumarins nor the β-carboline alkaloid demonstrated any toxic effect on digestive efficiencies of P. polyxenes in the presence of light; harmine, the alkaloid, did significantly reduce growth and consumption rates. None of the compounds had a significant effect on antioxidant enzyme levels. These findings contrast with those reported in earlier studies for Trichoplusia ni, a generalist noctuid sensitive to both furanocoumarins and β-carboline alkaloids. Greater detoxicative metabolic capabilities, coupled with substantially higher constitutive levels of antioxidant enzyme activity, likely explain at least in part the absence of induced antioxidant enzyme responses in the specialist feeder on phototoxic plants. © 1993 Wiley-Liss, Inc.     

The effect of host plant on larval survivorship of the Alaskan swallowtail butterfly (Papilio machaon aliaska)

The Alaskan swallowtail butterfly, Papilio machaon aliaska (Lepidoptera: Papilionidae), uses three plant species as host plants. Cnidium cnidiifolium (Turcz.) Schischk. belongs to the family Apiaceae, the ancestral host‐plant family of the P. machaon group. Artemisia arctica Less. and Petasites frigidus (L.) Franch, in contrast, belong to the distantly related Asteraceae family and were colonized relatively recently by this group of butterflies. Previous work has shown that larval survival is highest on the novel host plants when natural enemies are present in the field. Here I examine whether P. m. aliaska fitness varied when individuals were reared on the three host plants in a common environment, free of environmental and ecological complications such as predation. I collected 12 P. m. aliaska females and measured their reproductive success when their progeny were reared on each of the three host plants. I assessed larval fitness on each of the hosts by recording the percentage pupation and percentage emergence as well as by measuring pupal mass. I found that larvae reared on the ancestral host plant, C. cnidiifolium, had higher fitness than did larvae reared on either of the novel host plants, A. arctica or P. frigidus. Larvae reared on C. cnidiifolium were more likely to pupate, achieved a greater pupal mass, and had higher emergence rates than larvae reared on the novel hosts. I interpret these results to mean that, in the absence of predation, the ancestral host plant is a better host for P. m. aliaska larvae than either novel host and I contend that this result does not appear to support the hypothesis that P. m. aliaska populations in central Alaska are divided into host races.     

Poly modal emergence of the tiger swallowtail, Papilio glaucus (Lepidoptera: Papilionidae): source of a false second generation in central New York State

Abstract. 1. The tiger swallowtail butterfly, Papilio glaucus (L.) (Lepidoptera: Papilionidae), is commonly assumed to have a bivoltine life cycle in central New York State and other parts of the northeastern United States. The seasonal abundance of adults in Tompkins County, New York, shows a bimodal pattern in some years, and a skewed unimodal pattern in others. We studied the incidence of diapause, the potential for successful development of two generations in the field, and the dates of adult emergence from overwintering pupae to determine whether these seasonal patterns are the result of bivoltinism.
2. Insects from Tompkins County, reared as larvae under 16L:8D photoperiod, uniformly enter pupal diapause.
3. Comparison of the heat units required for successful development of two generations with the heat units available in the field during 11 years shows that emergence of non-diapausing individuals would occur too late in the summer to contribute to the second peak of butterfly abundance. Offspring of non-diapausing individuals could only rarely complete development before the end of the growth season.
4. Diapausing offspring of butterflies collected in early June and late July differed sufficiently in dates of emergence to account for the field pattern of seasonal abundance.
5. Electrophoretic studies revealed no evidence of genetic isolation between early and late emerging butterflies.
6. These results suggest that polymodal emergence of diapausing insects can masquerade as a bivoltine life cycle.     

Variation in phenology and nutritional quality between host plants and its effect on larval performance in a specialist butterfly,Zerynthia rumina

The main host plants of the butterfly Zerynthia rumina L. (Lepidoptera: Papilionidae) in southern Spain occur in different habitats and in general do not grow sympatrically. Therefore, each single local butterfly population uses the particular host available within its range. Aristolochia longa L. is a tuberous perennial herb available only in the spring, while A. baetica L. is an evergreen perennial vine with indeterminate growth. However, because of the toughness of older leaves, newly hatched larvae feed only on new leaves of A. baetica, and most of these leaves are produced well before the larvae hatch. In laboratory experiments, caterpillars feeding on either new or mature A. longa leaves grew faster and converted food into biomass more efficiently than those feeding on new A. baetica leaves. These differences are related to variation in nutritional quality among the host plants. Estimates of butterfly abundance were lower in sites where Z. rumina uses A. baetica, compared with those where the host is A. longa. The potential differential effect of these two food plants on the densities of local butterfly populations relying on them is discussed here.     

The relationship between feeding specialization and host plants to aldrin epoxidase activities of midgut homogenates in larval Lepidoptera

Abstract. 1. Aldrin epoxidase activities of midgut homogenates in fifty-eight species of phytophagous lepidopterous larvae, were determined.
2. There were no significant differences in mean activities amongst monophagous, oligophagous and polyphagous feeders. These results question the widely held view that there is a correlation between mixed function oxidase activity and feeding specialization.
3. Aldrin epoxidase activity depended largely on the host which was consumed. Larvae with high activity invariably fed on hosts containing monoterpene compounds.
4. The proposal that mixed function oxidases provide polyphagous insects with an effective biochemical defence against diverse allelochemicals is discussed.     

Segregation of F-2 interspecific hybrid growth performance and wing color patterns relative to parental species in the Papilio machaon species group （Lepidoptera： Papilionidae）

A strange Papilio female specimen with a prominent yellow wing band type very much like P. brevicauda （and also seen in P. machaon, and P. zelicaon） was captured in the forested mountains of southwest Vermont, USA. Despite its brevicauda-like appearance, it had mt-DNA of P. polyxenes. In eastern North America the P. brevicauda （Maritime Provinces of Canada） is currently believed to reflect previous （ancient） hybridization between ancestral P. machaon and P. polyxenes. It is uncertain whether the brevicauda-like wing trait features in the odd Vermont female may be a result of some introgression of P. machaon or P. brevicauda with P. polyxenes in eastern North America as seen also in the Ozark Mountains with P. joanae. Nonetheless, the significance of wing trait analyses in relation to hybrid zones with P. polyxenes remains relevant. The segregation of wing color patterns and hostplant use abilities （efficiencies and growth rates） were examined for laboratory hand-paired F-2 hybrids of P. zelicaon ~ Papilio polyxenes relative to the parental species and primary hybrids. The black adult color pattern characterizing P. polyxenes is presumed to be inherited as a simple autosomal dominant. These hybrids were examined to determine： （i） how F-1 primary hybrid offspring perform relative to the parental types in survival, growth rates, and growth efficiencies; and （ii） whether these larval traits and the adult color patterns might be linked with wing morphology in the F-2 hybrids. While the F-1 hybrids grew as well as the P. polyxenes parent （P. zelicaon was significantly less efficient and slower in growth than either of these）, we could not convincingly determine whether the growth performances of the F-2 segregants were linked with the parental wing patterns.     

Habitat use of the endangered butterfly Euphydryas maturna and forestry in Central Europe

The knowledge of ecological requirements of declining butterflies of European woodlands remains limited, which hinders conservation management of their localities. This also applies for continentally threatened scarce fritillary Euphydryas maturna . On the basis of the largest data set on its habitat use ever collected in Central Europe, we analyse habitat requirements of its populations in Austria (A), the Czech Republic (Cz) and Germany (D). All studied populations inhabit open-canopy sites within woodlands, but larval survival decreases under full sun and preferred sites are relatively humid and sheltered. Nests of pre-hibernation larvae occur at terminal branches of Fraxinus excelsior , 1.5–3 m above the ground. Pre-hibernation mortality reaches 70% (Cz, D). Another limiting factor is quality of woodland vegetation: post-hibernation larvae consume a wide range of herbs and shrubs, and adult distribution is linked to nectar availability. The butterfly thus depends on highly heterogeneous early successional stages of deciduous woods, historically maintained by coppicing (Cz, D) and forest pasture (A). Restoration of these traditional methods offers the only chance for survival of E. maturna in Central Europe, and the butterfly may become a flagship for other threatened organisms of open-canopy woodlands.     

Characteristics of larvae of the southwestern corn borer, Diatraea grandiosella, obtained from populations present in tropical and temperate regions

A comparative study was undertaken of southwestern corn borers (Diatraea grandiosella) collected from south central Mexico (19°N latitude) and southeast Missouri (37°N latitude). All the life stages of the Mexican insects were found to be larger or heavier, or both, than were those of the Missouri insects. Mexican larvae grew at a higher rate and attained a significantly heavier body weight than did Missouri larvae. Although both Mexican and Missouri larvae underwent ecdyses during diapause, Mexican larvae ecdysed more frequently than did the Missouri larvae at 23°C light:dark 12 h:12 h. Larvae that ecdysed most frequently did not necessarily spend longer in diapause. Electrophoresis of the fat body and haemolymph proteins of Mexican and Missouri larvae revealed similar patterns. The fat body of diapausing Mexican larvae contained substantial amounts of a diapause-associated protein which has been characterized previously from the fat body of Missouri Larvae. Double immunodiffusion confirmed that the diapause-associated protein of the Mexican larvae was identical to that present in the Missouri larvae. Smaller amounts of this protein appear to be present in the fat body of diapausing Mexican larvae than are present in that of diapausing Missouri larvae.     

Inter- and intra-specific differences in the number of larval instars in British populations of 24 species of stoneflies (Plecoptera)

1. Ontogenetic changes during the life cycle of aquatic insects are important not only in life‐history studies but also in evaluating food‐web structure. They require information on the growth and number of larval instars but such information is lacking for many species, including Plecoptera. Therefore, the chief objectives of the present study were to determine inter‐ and intra‐specific differences in the number of larval instars in British populations of 24 species of stoneflies, to test Dyar’s hypothesis that growth followed a geometric progression, and to synthesise this information with previously published values for four British species. 2. Larvae were reared at constant temperatures in the laboratory from eggs from 63 populations (one to six populations per species). First instars from each population were divided into three batches and each batch was reared at one of three constant temperatures. For each species, the rearing temperature and source population had no significant effect on the mean size of each larval instar. 3. The relationship between the geometric mean length of each instar and instar number was well described by an exponential equation (P < 0.001, r² > 0.9 for all species), thus supporting Dyar’s hypothesis. Only one species, Brachyptera risi, had the same number of instars for males and females (12–13). For the other 15 herbivorous species and the four smaller carnivorous ones, the number of instars was higher for females than males (range 11–16 for males, 12–17 for females). The larger size of the females was due to their additional instars, not a sex difference in growth rates. In contrast, there was a clear growth separation of the sexes after the 9th or 10th instar for the four largest carnivores. The number of larval instars was highest for these four species (range 16–19 for males, 18–23 for females), and females were much larger than males. 4. A multiple regression equation with data from the present and previous studies (n = 27) showed that variability in the mean length of the first instar and the maximum number of larval instars for each species accounted for 88% and 91% of the variability in the mean length of the final instar for males and females, respectively. 5. Values for Plecoptera in other countries were in general agreement with those in the present study, especially in the same families. Two old, but widely quoted, high values are doubtful. The present study and four previous ones provide a sound basis for ontogenetic studies on 28 species of Plecoptera and their role in aquatic ecosystems.     

Influence of plant sulphur nutrition on oviposition and larval performance of the diamondback moth

We measured adult oviposition preference, larval growth, and feeding behaviour of the crucifer specialist Plutella xylostella (L.) (Lepidoptera: Plutellidae) on plants of Brassica napus (L.) cv. Express (Brassicaceae), grown under three different sulphur regimes. The nutrient solutions used were the following: one sulphur‐free (S₀), one normal sulphur (S_n, normal field concentration), and one sulphur‐rich (S₊, double concentration of S_n). Females laid more eggs on S_n than on S₀ plants, while only a slight, non‐significant difference was observed between S_n and S₊ plants. Moreover, the development time from hatching to emergence was significantly shorter, and adults were heavier on S_n than on S₀ plants. Comparing these same two parameters from S_n and S₊ plants, we found a shorter development time on plants rich in sulphur, although this trend was not statistically significant. Larval feeding preferences were tested in a dual choice assay using leaf discs. A significantly higher number of larvae preferred leaf discs of S_n plants than those of S₀ plants. Furthermore, the larvae preferred S₊ to S_n discs. An optimal supply of sulphur to oilseed rape is necessary for a good seed harvest, and it also plays an important role in acceptance by P. xylostella of the host plant. Maintaining higher levels of sulphur in the plant nutrient solution benefits insect performance, both at the adult and larval stage.     

Trichome production and variation in young plant resistance to the specialist insect herbivore Plutella xylostella among natural populations of Arabidopsis lyrata

The strength of plant‐herbivore interactions varies spatially and through plant ontogeny, which may result in variable selection on plant defense, both among populations and life‐history stages. To test whether populations have diverged in herbivore resistance at an early plant stage, we quantified oviposition preference and larval feeding by Plutella xylostella (L.) (Lepidoptera: Plutellidae) on young (5–6 weeks old) Arabidopsis lyrata (L.) O'Kane & Al‐Shehbaz (Brassicaceae) plants, originating from 12 natural populations, six from Sweden and six from Norway. Arabidopsis lyrata can be trichome‐producing or glabrous, with glabrous plants usually receiving more damage from insect herbivores in natural populations. We used the six populations polymorphic for trichome production to test whether resistance against P. xylostella differs between the glabrous and the trichome‐producing morph among young plants. There was considerable variation among populations in the number of eggs received and the proportion of leaf area consumed by P. xylostella, but not between regions (Sweden vs. Norway) or trichome morphs. Rosette size explained a significant portion of the variation in oviposition and larval feeding. The results demonstrate that among‐population variation in resistance to insect herbivory can be detected among very young individuals of the perennial herb A. lyrata. They further suggest that trichome densities are too low at this plant developmental stage to contribute to resistance, and that the observed among‐population variation in resistance is related to differences in other plant traits.     

Differences in pupal cold hardiness and larval food consumption between overwintering and non‐overwintering generations of the common yellow swallowtail,Papilio machaon (Lepidoptera: Papilionidae), from the Osaka population

To clarify differences in pupal cold hardiness and larval food consumption between overwintering and non‐overwintering generations of the common yellow swallowtail, Papilio machaon, we reared larvae from the Osaka population under photoperiods of 16 h light : 8 h dark (LD 16:8) (long day) or LD 12:12 (short day) at 20°C. We examined the relationship between food consumption and weight during the final larval stadium and pupae, and measured the pupal supercooling point (SCP). Although the ratio of assimilation to consumption did not differ significantly between photoperiods, the ratio of assimilation to pupal weight differed significantly between individuals reared under long and short days. All diapausing pupae were brown, whereas 56% of non‐diapausing pupae were green with the remainder brown. The mean pupal body length (L), dorsal width (W₁) and lateral width (W₂) were larger in non‐diapausing than in diapausing pupae, and the W₁/L and W₁/W₂ ratios differed significantly between non‐diapausing and diapausing pupae. SCP was approximately –20°C and did not differ among pupae 5, 15 and 30 days after pupation under long‐day conditions. However, under short‐day conditions, mean SCP gradually decreased, stabilizing at approximately –24 to –25°C by 30 days after pupation. After freezing, some diapausing pupae emerged as adults, whereas all non‐diapausing pupae died. Both egestion and assimilation were greater under long‐day conditions. The results revealed that pupae of this papilionid exhibit seasonal polyphenism in physiological and morphological traits. Energy from food appears to be expended on increasing cold hardiness in the overwintering generation and on reproduction in the non‐overwintering generation.     

Correlated responses to selection for faster development and early reproduction in Drosophila: the evolution of larval traits

Studies on selection for faster development in Drosophila have typically focused on the trade-offs among development time, adult weight, and adult life span. Relatively less attention has been paid to the evolution of preadult life stages and behaviors in response to such selection. We have earlier reported that four laboratory populations of D. melanogaster selected for faster development and early reproduction, relative to control populations, showed considerably reduced preadult development time and survivorship, dry weight at eclosion, and larval growth rates. Here we study the larval phase of these populations in greater detail. We show here that the reduction in development time after about 50 generations of selection is due to reduced duration of the first and third larval instars and the pupal stage, whereas the duration of the second larval instar has not changed. About 90% of the preadult mortality in the selected populations is due to larval mortality. The third instar larvae, pupae, and freshly eclosed adults of the selected populations weigh significantly less than controls, and this difference appears during the third larval instar. Thereafter, percentage weight loss during the pupal stage does not differ between selected and control populations. The minimum amount of time a larva must feed to subsequently complete development is lower in the selected populations, which also exhibit a syndrome of reduced energy expenditure through reduction in larval feeding rate, larval digging and foraging activity, and pupation height. Comparison of these results with those observed earlier in populations selected for adaptation to larval crowding and faster development under a different protocol from ours reveal differences in the evolved traits that suggest that the responses to selection for faster development are greatly affected by the larval density at which selection acts and on details of the selection pressures acting on the timing of reproduction.     

Temperature‐ and sex‐related effects of serine protease alleles on larval development in the Glanville fritillary butterfly

The body reserves of adult Lepidoptera are accumulated during larval development. In the Glanville fritillary butterfly, larger body size increases female fecundity, but in males fast larval development and early eclosion, rather than large body size, increase mating success and hence fitness. Larval growth rate is highly heritable, but genetic variation associated with larval development is largely unknown. By comparing the Glanville fritillary population living in the Åland Islands in northern Europe with a population in Nantaizi in China, within the source of the post‐glacial range expansion, we identified candidate genes with reduced variation in Åland, potentially affected by selection under cooler climatic conditions than in Nantaizi. We conducted an association study of larval growth traits by genotyping the extremes of phenotypic trait distributions for 23 SNPs in 10 genes. Three genes in clip‐domain serine protease family were associated with larval growth rate, development time and pupal weight. Additive effects of two SNPs in the prophenoloxidase‐activating proteinase‐3 (ProPO3) gene, related to melanization, showed elevated growth rate in high temperature but reduced growth rate in moderate temperature. The allelic effects of the vitellin‐degrading protease precursor gene on development time were opposite in the two sexes, one genotype being associated with long development time and heavy larvae in females but short development time in males. Sexually antagonistic selection is here evident in spite of sexual size dimorphism.     

Genetic analysis of diapause capability and association between larval and pupal photoperiodic responses in the flesh fly Sarcophaga similis

Abstract A variant of the flesh fly Sarcophaga similis, which does not enter pupal diapause even under diapause‐inducing conditions, is established by artificial selection. Interline crosses of the wild‐type and the variant revealed that the diapause capability is inherited in an incomplete dominant manner. Neither sex linkage nor a maternal factor is involved in the mode of inheritance. Genetic and genetic–environmental interactions are involved in the induction of diapause. In the wild‐type, the post‐feeding larval period is prolonged in response to short days. However, in the variant, the duration of the larval period under short‐day conditions is identical to that under long‐day conditions. Long‐day responses under short‐day conditions at both the larval and pupal stages in the variant indicate that the responses are established by common factors in both stages. The factors causing the long‐day responses under short‐day conditions at different developmental stages in the variant remain to be identified. However, it is plausible to hypothesize that common factors regulate ecdysteroid release at both the larval and pupal stages, and that malfunction of the system regulating these factors triggers such ecdysteroid release, irrespective of the photoperiod. Hence, the photoperiodic responses disappear in the variant in both stages. The adult stage of the variant has a functional circadian clock, suggesting that the nondiapause phenotype is not necessarily involved in malfunction of the circadian clock.     

Geographic variation in growth and food conversion efficiency of juvenile Atlantic halibut related to latitude

Higher growth capacity and food conversion efficiency was observed in populations of juvenile halibut from high Hippoglossus hippoglossus compared lower latitudes. In addition, temperature adaptation shown by the lower temperature optimum for growth in the Norwegian population (mean±S.E. 12·9±0·10 C) compared with the Icelandic and Canadian populations (14·2±0·2 and 13·9±0±30 C respectively), seems to occur. Overall the data support the hypothesis of countergradient variation in growth. These results have implications firstly for selection focusing on growth performance in halibut culture; and secondly, for safe prediction of growth, since if countergradient variation in growth performance occurs one cannot assume automatically that a species will respond to the same set of physiological parameters in the same way throughout its range.     

Differentiation of roles of chemosensory organs in food discrimination among host and non-host plants by larvae of the tobacco hornworm, Manduca sexta

ABSTRACT. The contributions of olfactory and gustatory organs in food plant discrimination were examined in larvae of Manduca sexta (Johan.) (Lepidoptera, Sphingidae). Larvae, from which various chemosensory organs had been removed surgically, were tested for feeding preferences for a host, tomato ( Lycopersicon esculentum ); a weakly acceptable non-host, rape ( Brassica napus ); and an unacceptable non-host canna ( Canna generalis ), using a two-choice disc bioassay.
Removal of all known chemosensory organs resulted in failure to show discriminatory behaviour in a strictly chemosensory bioassay, indicating that all external chemosensory organs have been accounted for. The involvement of non-chemosensory organs results in residual discrimination for leaves by individuals with total chemosensory ablations.
Larvae possessing either olfactory or gustatory organs still exhibit normal preferences for tomato over rape. Gustatory (but not olfactory) organs are required for larvae to show normal preferences for tomato over canna; in fact, olfactory organs do not appear to participate in this decision.
To examine which if any of the plant species is being selected in two-choice tests, larvae were given a choice between each leaf species and a 'neutral' substance (wet filter paper). Both olfactory and gustatory organs are required for normal preferences for tomato, but either alone will suffice for rape. Only gustation is needed to select canna, and participation of either the epipharyngeal sensilla or a single medial sensillum styloconicum is sufficient to elicit complete rejection behaviour.
We conclude that, in larvae of M. sexta , the complement of chemosensory organs needed for food plant discrimination varies with the plant species sampled. Evidence is presented exposing a potential artefact of ablation experiments; extirpation of one sensory organ may affect the functioning of others nearby, even though they may appear normal by visual inspection.