Agonistic display or courtship behavior? A review of contests over mating opportunity in butterflies

Male butterflies compete over mating opportunities. Two types of contest behavior are reported. Males of various butterfly species compete over a mating territory via aerial interactions until one of the two contestants retreats. Males of other butterfly species fly around larval food plants to find receptive females. Males of some species among the latter type can find a conspecific pupa, and they gather around it without expelling their rivals. Scramble competition over mating occurs when a female emerges from the pupa. Many studies have been performed on territorial species, and their contest resolution has often been understood from the point of view of contest models based on game theory. However, these models cannot explain why these butterflies perform contest displays despite the fact that they do not have the ability to attack their opponent. A recent study based on Lloyd Morgan’s Canon showed that territorial contests of male butterflies are better understood as erroneous courtship between sexually active males. In this paper, I review research on contests over mating opportunity in butterflies, and show that the erroneous courtship framework can explain not only territorial contests of butterflies but also why males do not determine the owner of a conspecific pupa.     

Are tropical butterflies more colorful?

There is a common and long-standing belief that tropical butterflies are more striking in their coloration than those of cooler climates. It has been suggested that this is due to more intense biotic selection or mate selection in the tropics. We tested whether there were differences in coloration by examining the dorsal surface color properties of male butterflies from three regions of the western hemisphere: the Jatun-Satcha Reserve in lowland Ecuador (tropical), the state of Florida, USA (subtropical) and the state of Maine, USA (cool temperate). We digitally photographed the dorsal wing and body surface of male butterfly specimens from Maine, Florida, and Ecuador. For each photograph, we analyzed the mean and variation for the color-parameters that are thought to be related to colorfulness; namely Hue, saturation and intensity. Overall, the Ecuadorian sample exhibited more varied intensity, saturation, and Hue compared to the other regions. These results suggest a more complex assemblage of colors and patterns regionally and on a butterfly-by-butterfly basis in the tropics. The greater complexity of colors within each butterfly in our Ecuadorian sample suggests that tropical butterflies are indeed more ‘colorful’, at least by some measures. Possible reasons for this include stronger predation pressure selecting for aposematism, greater species diversity selecting for camouflage or warning coloration against potential predators, and easier recognition of potential mates in a species rich environment.     

Contest behavior in territorial male butterflies: does size matter?

Although intrasexual contests generally favor bigger or stronger individuals, the relevance of body size to war of attrition-typedisputes between weaponless animals such as butterflies isunclear. In this study I aimed to investigate the significanceof size in this context by studying territorial contests inHypolimnas bolina (L.), a species that exhibits consistentseasonal plasticity in body size. In this species adult ageis positively correlated with large size in spring but withsmall size in autumn. This shift allowed independent evaluationof the relevance of each variable (size and age) to intrasexualcontest success. Observation of a population of marked individualsindicated that only age appeared important, with the winnersof pair-wise contests significantly older than losers in both seasons, and with contests lasting longer when the age differencebetween the combatants was small. Age was also linked to residency;residents won 99% of all contests. This study suggests thatsize does not matter in these aerial disputes, but age andresidency do. It is not yet possible to determine whether olderbutterflies are intrinsically better competitors, or whether they simply have greater opportunity to find a vacant territory.     

A search method for homologs of small proteins. Ubiquitin-like proteins in prokaryotic cells?

The question of protein homology versus analogy arises when proteins share a common function or a common structural fold without any statistically significant amino acid sequence similarity. Even though two or more proteins do not have similar sequences but share a common fold and the same or closely related function, they are assumed to be homologs, descendant from a common ancestor. The problem of homolog identification is compounded in the case of proteins of 100 or less amino acids. This is due to a limited number of basic single domain folds and to a likelihood of identifying by chance sequence similarity. The latter arises from two conditions: first, any search of the currently very large protein database is likely to identify short regions of chance match; secondly, a direct sequence comparison among a small set of short proteins sharing a similar fold can detect many similar patterns of hydrophobicity even if proteins do not descend from a common ancestor. In an effort to identify distant homologs of the many ubiquitin proteins, we have developed a combined structure and sequence similarity approach that attempts to overcome the above limitations of homolog identification. This approach results in the identification of 90 probable ubiquitin-related proteins, including examples from the two prokaryotic domains of life, Archaea and Bacteria.     

Is male puddling behaviour of tropical butterflies targeted at sodium for nuptial gifts or activity?

An apparent sexual difference in adult feeding behaviour in many species of Lepidoptera relates to puddling on mud, dung and carrion. In most butterfly species, puddling is exclusively a male behaviour. A possible explanation for this division in feeding behaviour is that nutrients derived from puddling are transferred to the female in the spermatophore during mating as a nuptial gift. Sodium derived from puddling has been shown to act as a nuptial gift in a few Lepidoptera species. It can also be used for neuromuscular activity in both males and females and may therefore correlate with flight morphology. In this study, we examine the generality of these two hypotheses in comparative work on a community of African fruit-feeding butterflies. We investigated puddling behaviour of males and females on carrion and dung together with sodium preferences, polyandry, relative wing-size, sexual size dimorphism and sodium concentrations in the bodies and spermatophores of several species. The results show that sodium as a nuptial gift can explain the sexual division in puddling in some species, but not in all. Species in which both sexes puddle transfer little sodium in the nuptial gift, which is consistent with the nuptial gift theory. Wing loading and puddling are not significantly correlated, but the trend followed the direction predicted by the activity hypothesis. However, the sodium concentration in the species with the smallest wing area to thoracic volume (WA/TV) ratio (the largest Charaxes spp.), was relatively low. Moreover, in all investigated species, the sodium concentration was higher in the abdomen than in the thorax. The results are discussed in the light of differences between the sexes in foraging behaviour in both larvae and adults, and with respect to alternative explanations for puddling.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 345–361.     

Carbamylation of proteins in 2-D electrophoresis--myth or reality?

Carbamylation is widely quoted as being a problem in 2-D gel analysis and the associated sample preparation steps. This modification occurs when iso-cyanate, a urea break-down product, covalently modifies lysine residues, thus inducing a change in isoelectric point. Urea is used at up to 9 M concentrations in sample preparation and 2-D gels because of its ability to disrupt protein structure and effect denaturation without the need for ionic surfactants such as SDS. We have studied carbamylation using 7 M urea and 2 M thiourea, under a range of experimental temperatures to establish when, and if, it occurs and what can be done to minimize the modification. The actual time required for protein extraction from a tissue is usually short compared to the time required for procedures such as reduction and alkylation and IPG rehydration and focusing. Therefore, it is the temperature during these post-extraction procedures that is the most critical factor. Our experiments have shown that carbamylation does not occur during electrophoresis in the presence of urea, even with prolonged run-times. However, under poorly controlled sample preparation and storage conditions, it can become a major event.     

Diversification of host use in two polyphagous butterflies: differences in oviposition specificity or host rank hierarchy?

Novel host usage may represent an initial step towards diversification or radiation onto novel hosts within an evolutionary lineage, particularly if a shift in host plant preference ranking takes place. Polyphagous stages of evolutionary lineages may represent transitional states in which novel host associations are more likely to develop, but may be more difficult to detect experimentally. The polyphagous sister species Papilio glaucus L. and Papilio canadensis (Lepidoptera: Papilionidae; these Papilio = Pterourus) are known to exhibit differences in host‐plant use, despite significant overlap in host‐use abilities, providing an opportunity to examine how host shifts in polyphagous species may occur and what the implications for future divergence may be. In particular, we were interested in (i) determining whether differences in oviposition behavior of these species were due to changes in specificity or shifts in host‐plant hierarchy, (ii) whether the varying preference for primary hosts also affected the preference for secondary hosts, and (iii) what the oviposition preferences of a new hybrid swarm population are. We examined more than 40 000 oviposition bouts from more than 400 P. glaucus, P. canadensis, and hybrid females placed in seven‐, three‐, or two‐choice assays. In each of the choice assays, leaves from plants in different plant families of varying suitability for P. glaucus and P. canadensis larvae were used. We found the primary difference between P. glaucus and P. canadensis to be limited to a Z‐linked shift in host rank hierarchy due to an acceptance of Populus tremuloides Michx. (Salicaceae) and reduced specificity for Liriodendron tulipifera L. (Magnoliaceae) in P. canadensis. In addition, we found the absence of the Z‐linked oviposition acceptance of P. tremuloides in a recently formed allochronically separated hybrid swarm population found in P. canadensis territory at the northern border of the P. glaucus and P. canadensis hybrid zone.     

Nucleocytoplasmic trafficking of proteins: With or without Ran?

Proteins and RNAs move between the nucleus and cytoplasm by translocation through nuclear pore complexes in the nuclear envelope. To do this, they require specific targeting signals, energy, and a cellular apparatus that catalyzes their transport. Several of the factors involved in nucleocytoplasmic trafficking of proteins have been identified and characterized in some detail. The emerging picture for nuclear transport proposes a central role for the small GTPase Ran and proteins with which it interacts. In particular, asymmetric distribution of these proteins between nucleus and cytoplasm appears to be responsible for the vectorial nature of nucleocytoplasmic transport. Here, we summarize the role of Ran and Ran-binding proteins in nuclear trafficking of proteins with classical nuclear localisation signals. We also discuss examples of the growing number of alternative pathways that are involved in transport of proteins across the nuclear envelope. BioEssays 21:579–589, 1999. © 1999 John Wiley & Sons, Inc.     

Six authors in search of a citation: villains or victims of the Vancouver convention?

OBJECTIVES--To analyse trends in the number of authors per article over the past 10 years. DESIGN--Analysis of articles from random volumes of eight biomedical journals. SUBJECTS--Cell, Nature, Proceedings of the National Academy of Sciences USA (PNAS), Journal of Clinical Investigation (JCI), Biochemical and Biophysical Research Communications (BBRC), Journal of Clinical Oncology (JCO), New England Journal of Medicine (NEJM), Lancet. MAIN OUTCOME MEASURES--Median and modal numbers of authors. RESULTS--All journals except Cell and Nature showed a trend towards increasing authorship numbers over the study period. The trend was most noticeable in journals such as JCO which feature clinical research. General medical journals (Lancet, NEJM) with a median of six to seven authors per article published far fewer seven author than six author studies, which suggests that author number may be influenced by the Vancouver convention which precludes citation of more than six authors. CONCLUSIONS--The phenomenon of expanding authorship in biomedical journal articles is not explained by the hypothesis that newer research technologies have necessitated more extensive collaboration. Rather, the data suggest that conferral of authorship may sometimes have a volitional component which contributes to rising author numbers. It is proposed that replacement of the Vancouver convention with a "first author, last author" citation system may help stem this rise in author numbers.     

Bcl-2 proteins: regulators of apoptosis or of mitochondrial homeostasis?

Programmed cell death (apoptosis) is used by multicellular organisms during development and to maintain homeostasis within mature tissues. One of the first genes shown to regulate apoptosis was bcl-2. Subsequently, a number of Bcl-2-related proteins have been identified. Despite overwhelming evidence that Bcl-2 proteins are evolutionarily conserved regulators of apoptosis, their precise biochemical function remains controversial. Three biochemical properties of Bcl-2 proteins have been identified: their ability to localize constitutively and/or inducibly to the outer mitochondrial, outer nuclear and endoplasmic reticular membranes, their ability to form heterodimers with proteins bearing an amphipathic helical BH3 domain, and their ability to form ion-conducting channels in synthetic membranes. The discovery that mitochondria can play a key part in the induction of apoptosis has focused attention on the role that Bcl-2 proteins may have in regulating either mitochondrial physiology or mitochondria-dependent caspase activation. Here we attempt to synthesize our current understanding of the part played by mitochondria in apoptosis with a consideration of how Bcl-2 proteins might control cell death through an ability to regulate mitochondrial physiology.     

Convergence and extension in vertebrate gastrulae: cell movements according to or in search of identity?

During vertebrate gastrulation, convergence and extension cell movements both narrow and lengthen the forming embryonic axis. Concurrently, positional information established principally by the ventral-to-dorsal gradient of bone morphogenetic protein activity specifies cell fates within the gastrula. New data, primarily from zebrafish, have identified domains of distinct convergence and extension movements, and have established a role for the noncanonical Wnt signaling pathway in promoting the mediolateral cell polarization that underlies this morphogenetic process. Other observations suggest the intriguing possibility that positional information regulates convergence and extension movements in parallel with cell-fate specification.     

Kasha or state selective behavior in the photochemistry of ortho-nitrobenzaldehyde?

The photochemistry of ortho-nitrobenzaldehyde dissolved in tetrahydrofuran was studied by means of femtosecond UV/Vis and IR spectroscopy. Comparison was made of the spectral and temporal signatures for ~400 nm and ~260 nm excitation. The 400 nm excitation promotes NBA to its lowest excited singlet state of nπ* character whereas for 260 nm an upper excited state of ππ* character is addressed. On the picosecond time scale, the molecule undergoes hydrogen transfer, yielding a ketene intermediate, internal conversion recovering the starting material, and intersystem crossing. Time constants and yields of these processes are virtually not affected by the excitation wavelength. For 400 nm excitation a ~100 fs decay component seen in the 260 nm experiment is absent, indicating that this component is due to a ππ* → nπ* internal conversion. In contrast to its formation, the decay of the ketene intermediate is influenced by the excitation wavelength. This can be attributed to different amounts of vibrational excitation.     

Misfolding of membrane proteins in health and disease: the lady or the tiger?

Protein misfolding is increasingly recognized as a factor in many diseases, including cystic fibrosis, Parkinson's, Alzheimer's and atherosclerosis. Many proteins involved in misfolding-based pathologies are membrane-associated, such that the bilayer may play roles in normal and aberrant folding. It can be argued that the in vivo partitioning of eukaryotic membrane proteins between folding and misfolding pathways is under kinetic control. Moreover, the balance between these pathways can be surprisingly delicate.     

Xylem-borne cytokinins: still in search of a role?

Leaf expansion and xylem cytokinin concentration ([X-CK]) decrease in response to nitrogen (N) deprivation. Debate continues over cause, effect, and correlation. Supporting studies provide, at best, correlative evidence that [X-CK] controls leaf growth in response to N-deprivation, while dissenting studies indicate that leaf growth responses to N can be independent of changes in X-CK supply to leaves. A model is proposed to evaluate the physiological significance to leaf growth of changes in plant and environment N concentrations, and plant CK concentrations.     

Plakophilins: Multifunctional proteins or just regulators of desmosomal adhesion?

Plakophilins 1-3 are members of the p120(ctn) family of armadillo-related proteins. The plakophilins have been characterized as desmosomal proteins, whereas p120(ctn) and the closely related delta-catenin, ARVCF and p0071 associate with adherens junctions and play essential roles in stabilizing cadherin mediated adhesion. Recent evidence suggests that plakophilins are essential components of the desmosomal plaque where they interact with desmosomal cadherins as well as the cytoskeletal linker protein desmoplakin. Plakophilins stabilize desmosomal proteins at the plasma membrane and therefore may function in a manner similar to p120(ctn) in the adherens junctions. The three plakophilins reveal distinct expression patterns, and although partially redundant in their function, mediate distinct effects on desmosomal adhesion. Besides a structural role, a function in signaling has been postulated in analogy to other armadillo proteins such as beta-catenin. At least plakophilins 1 and 2 are also localized in the nucleus, and all three proteins occur in a cytoplasmic pool. This review aims to summarize the current knowledge of plakophilin function in the context of cell adhesion, signaling and their putative role in diseases.     

Ipomoea bahiensis pollinators: Bees or butterflies?

Ipomoea bahiensis presents many characters of a psychophilous nectar flower. At the investigated synanthropic site, it is, however, visited and pollinated primarily by pollen-foraging small to medium sized bees that leave the flowers well before the end of anthesis. The busy bees appear to intimidate, probably by their mere existence, butterflies that function as pollinators in early morning, but disappear from the scene after beginning of anthesis. The plant is allogamous. A higher fruit set in hand cross-pollinated flowers as compared to naturally pollinated controls indicates pollen limitation. For the first time, we observed male bees entering the closing flowers as an overnight accommodation even before noon.     

Diet divergence in two sympatric congeneric butterflies: Community or species level phenomenon?

Summary Two species ofEuphydryas butterflies were studied in California, USA, and showed considerable diet overlap at the species level. They utilize many of the same plant genera for oviposition. However,E. editha is less likely to use woody perennials than isE. chalcedona.Both butterfly species are known to specialize on different host plants in different populations, so species level divergence may not be a good predictor of community level divergence. Within five communities,E. editha andE. chalcedona showed no dietary overlap. A major component of the niche ofE. editha in one community was occupied byE. chalcedona in a second community, even though both butterfly species occupied both communities. These resource use patterns indicate that community level interactions may affect diet divergence. The degree to which divergence within communities is greater (or less) than expected from a species level comparison may be used to provide a measure of community organization. Equations are given in the Appendix for calculating overlap probabilities from presence/absence types of data; in this study, presence is oviposition on a particular plant species, absence is no oviposition on that plant species. Given the various assumptions of the model,E. editha andE. chalcedona show significant community level components of their dietary divergence in the areas studied. However, in some other communitiesE. editha andE. chalcedona do share host plant species. Therefore, we could not demonstrate community level divergence conclusively, nor has this been demonstrated for any other pair of insect herbivore species. We do not know whether this is because the phenomenon is truly rare or just very hard to demonstrate.