Lateralised courtship behaviour and its impact on mating success in Ostrinia furnacalis (Lepidoptera: Crambidae).

Lateralisation is a well-established phenomenon observed in an increasing number of insect species. This study aims to obtain basic details on lateralisation in courtship and mating behaviour in Ostrinia furnacalis, the Asian corn borer. We conducted laboratory investigations to observe lateralisation in courtship and mating behaviours in adult O. furnacalis. Our goal was also to detect lateralised mating behaviour variations during sexual interactions and to elucidate how these variances might influence the mating success of males. Our findings reveal two distinct lateralised traits: male approaches from the right or left side of the female and the direction of male turning displays. Specifically, males approaching females from their right side predominantly exhibited left-biased 180° turning displays, while males approaching females from the left-side primarily displayed right-biased 180° turning displays. Notably, left-biased males, executing a 180° turn for end-to-end genital contact, initiated copulation with fewer attempts and began copulation earlier than their right-biased approaches with left-biased 180° turning displays. Furthermore, mating success was higher when males subsequently approached the right side of females during sexual encounters. Left-biased 180° turning males exhibited a higher number of successful mating interactions. These observations provide the first report on lateralisation in the reproductive behaviour of O. furnacalis under controlled laboratory conditions and hold promise for establishing reliable benchmarks for assessing and monitoring the quality of mass-produced individuals in pest control efforts.

Genomic regions underlying the species-specific mating songs of green lacewings.

Rapid species radiations provide insight into the process of speciation and diversification. The radiation of Chrysoperla carnea-group lacewings seems to be driven, at least in part, by their species-specific pre-mating vibrational duets. We associated genetic markers from across the genome with courtship song period in the offspring of a laboratory cross between Chrysoperla plorabunda and Chrysoperla adamsi, two species primarily differentiated by their mating songs. Two genomic regions were strongly associated with the song period phenotype. Large regions of chromosomes one and two were associated with song phenotype, as fewer recombination events occurred on these chromosomes relative to the other autosomes. Candidate genes were identified by functional annotation of proteins from the C. carnea reference genome. The majority of genes that are associated with vibrational courtship signals in other insects were found within QTL for lacewing song phenotype. Together these findings suggest that decreased recombination may be acting to keep together loci important to reproductive isolation between these species. Using wild-caught individuals from both species, we identified signals of genomic divergence across the genome. We identified several candidate genes both in song-associated regions and near divergence outliers including nonA, fruitless, paralytic, period, and doublesex. Together these findings bring us one step closer to identifying the genomic basis of a mating song trait critical to the maintenance of species boundaries in green lacewings.

Iridescent scales signal male fighting ability to access females in the jewelled splitfin Xenotoca variata.

Jewelled splitfin males (Xenotoca variata) possess multi-colour iridescent scales (speckles) on the flanks. This study tested the hypothesis that the number of speckles could be a good proxy for predicting fighting ability in contests for access to females. The experiments consisted in observing and recording males' agonistic behaviour, courtship displays and mating attempts in mixed-sex groups. The data were analysed and presented based on a Bayesian approach, which revealed that the density (cm-2 ) of speckles was positively correlated with the increase observed in the proportional frequency of attacks. Similarly, the density of speckles was positively associated with the increase observed in the frequency of courtship behaviour and with the frequency of mating attempts. Male-male aggressions drastically diminished (by eightfold) when females were removed from the observation tanks. These results indicate that the number of speckles is a consistent predictor of successful access to females and therefore, speckled males are likely to be able to gain more mating opportunities. This finding highlights the ubiquitous role iridescent colours play in visual signalling.

Beetle-robot hybrid interaction: sex, lateralization and mating experience modulate behavioural responses to robotic cues in the larger grain borer Prostephanus truncatus (Horn).

Ethorobotics, a new fascinating field of biorobotics, proposes the use of robotic replicas as an advanced method for investigating animal behaviour. This novel research approach can also encourage the development of advanced bioinspired robots. In the present study, we investigated the pushing behaviour, a particular display occurring in several beetle species, such as the larger grain borer, Prostephanus truncatus, during both male-female and male-male contexts. We developed a robotic apparatus actuating female and male-mimicking dummies to study if sex, mating experience and asymmetries of robotic cues can modulate the escalation of pushing behaviour. Results showed that the time needed by P. truncatus to react to female-smelling biomimetic dummies was chiefly affected by their mating experience and the dummy odour. This was likely due to reduce waste of costly sperm in mated males during the subsequent sexual interactions. The pushing behaviour was performed longer and with a higher number of acts when virgin females were approached from their right side. More and longer pushing acts were noted when virgin males were approached from their left side. Dedicated neural circuits would likely act in opposite direction in females and males producing population-level lateralized sensory-motor displays, which may be evolved to promote male approaches from the left side of females, thus improving short-distance sex recognition. Overall, this study provides new insights on the behavioural ecology of stored-product beetles, as well as on self-organization and decentralized decision making that can be exploited to develop bioinspired algorithms for task optimization, involving real-world scenarios.

Continuous exposure to mercury during embryogenesis and chick development affects later survival and reproduction of zebra finch (Taeniopygia guttata).

Methylmercury (MeHg) is a global environmental contaminant that bioaccumulates and has multiple toxic modes of action. Aquatic species have traditionally been the focus of wildlife toxicological research on mercury, but terrestrial organisms, including passerine birds, can be exposed to similarly elevated levels of MeHg. In this study we exposed a model passerine, the zebra finch (Taeniopygia guttata), to MeHg in ovo, as chicks only, or with a combined 'in ovo + chick' treatment. We isolated exposure to specific developmental stages through the use of egg injections (3.2 µg Hg/g egg) and controlled oral dosing of chicks (0.24 µg Hg/g bw/day from day 1 to day 30). In ovo exposure to MeHg reduced hatching success, but there was no effect of MeHg on chick growth. We found that in ovo only or chick only exposure did not have long-term effects, but there was some evidence for longer-term effects of combined 'in ovo + chick' exposure on post-fledging survival and potentially sex-biased survival which resulted in very few 'in ovo + chick' exposed females surviving to breed. These females also had lower overall breeding productivity that was mainly due to lower hatching success of their offspring, not lower chick-rearing success. We found no effect of treatment on clutch size or latency to laying among females that did lay eggs. Our study suggests that combined embryonic and nestling MeHg exposure has compounding latent effects on productivity, likely through a mechanism that influences the ability of females to lay fertile eggs that hatch.

Variation in sexual brain size dimorphism over the breeding cycle in the three-spined stickleback.

Snapshot analyses have demonstrated dramatic intraspecific variation in the degree of brain sexual size dimorphism (SSD). Although brain SSD is believed to be generated by the sex-specific cognitive demands of reproduction, the relative roles of developmental and population-specific contributions to variation in brain SSD remain little studied. Using a common garden experiment, we tested for sex-specific changes in brain anatomy over the breeding cycle in three-spined stickleback (Gasterosteus aculeatus) sampled from four locations in northern Europe. We found that the male brain increased in size (ca. 24%) significantly more than the female brain towards breeding, and that the resulting brain SSD was similar (ca. 20%) for all populations over the breeding cycle. Our findings support the notion that the stickleback brain is highly plastic and changes over the breeding cycle, especially in males, likely as an adaptive response to the cognitive demands of reproduction (e.g. nest construction and parental care). The results also provide evidence to suggest that breeding-related changes in brain size may be the reason for the widely varying estimates of brain SSD across studies of this species, cautioning against interpreting brain size measurements from a single time point as fixed/static.

A reply to Nieberding and Holveck: beyond experimental design and proximate mechanisms - mate choice in the face of sexual conflict.

We summarise our work on male mating behaviour in the tropical butterfly Bicyclus anynana, responding to the commentary provided by Nieberding and Holveck. We acknowledge that our laboratory studies are not free of shortcomings and potential caveats, though we attempted to address or highlight these within each paper. The concerns raised seem to stem mainly from different notions with respect to the proximate basis of old male mating advantage, and specifically the relative importance of male behaviour versus pheromone blend. In our view, our experiments provided compelling evidence for a prominent role of male behaviour, while we were unable to obtain clear evidence for a major role of male sexual pheromones. In addition to the lack of evidence we argue that a preference of females for older males based on pheromone blend is unlikely, as pheromone titres do not seem to indicate male quality and, more importantly, females actually suffer a fitness cost when mating with older males. The latter suggests that old male mating advantage arises from sexual conflict rather than cooperation. We thus highlight the importance of considering both the proximate and the ultimate level for gaining an integrative understanding of complex behavioural patterns.

Social context-dependent modification of courtship behaviour in Drosophila prolongata.

Induction of alternative mating tactics by surrounding conditions, such as the presence of conspecific males, is observed in many animal species. Satellite behaviour is a remarkable example in which parasitic males exploit the reproductive investment by other males. Despite the abundance of parasitic mating tactics, however, few examples are known in which males alter courtship behaviour as a counter tactic against parasitic rivals. The fruit fly Drosophila prolongata shows prominent sexual dimorphism in the forelegs. When courting females, males of D. prolongata perform 'leg vibration', in which a male vibrates the female's body with his enlarged forelegs. In this study, we found that leg vibration increased female receptivity, but it also raised a risk of interception of the female by rival males. Consequently, in the presence of rivals, males of D. prolongata shifted their courtship behaviour from leg vibration to 'rubbing', which was less vulnerable to interference by rival males. These results demonstrated that the males of D. prolongata adjust their courtship behaviour to circumvent the social context-dependent risk of leg vibration.

Colour Selection and Olfactory Responses of Papilio demoleus during Foraging and Courtship.

Colours and odours are the most important cues for butterflies to localise food and mating partners. We studied the visual and olfactory responses of the widely distributed butterfly Papilio demoleus Linnaeus during foraging and courtship. P. demoleus visited odourless flowers with six colours except green and black, with red as the favourite colour (650-780 nm). Males and females differed in behaviour while visiting flowers. Males were more active than females during foraging. The application of honey water resulted in a significant increase in flower visits by both females and males, and they scarcely visited the apetalous branches with odours. Under natural conditions, four patterns were observed: males chasing males (42.28%), males chasing females (30.56%), females chasing females (13.73%), and females chasing males (13.43%). Males chasing males was the most frequent, probably because males drive away competing con-specific males. When butterflies visited odourless mimics, males chasing females (70.73%) and males chasing males (29.27%) also occurred, indicating that males could accurately distinguish mates using colours only without any chemical cues, and females need chemical cues. The behavioural responses of P. demoleus to floral visits and courtship suggest that colour is the dominant factor during foraging and courtship. We verified the presence of P. demoleus rhodopsin genes, including Rh2, Rh3, Rh4, and Rh5, for long wavelength, blue, and ultraviolet (UV) spectrum recognition, which is consistent with the colour recognition of flowers and wings during visiting flowers and courtship.

Male-male associations in spotted bowerbirds (Ptilonorhynchus maculatus) exhibit attributes of courtship coalitions.

Abstract: Despite strong selective pressures inherent in competition for mates, in species with non-resource-based mating systems males commonly engage in non-agonistic interactions with same-sex visitors at display arenas. Bowerbirds perform courtship dances on elaborate display structures - known as bowers - that are built and defended by one resident male. Several reports have suggested that bower owners tolerate the presence of specific male visitors at their display arenas, referred to here as 'subordinates'. Subordinate males may learn the skills required for successful sexual signalling via prolonged social interactions at adults' arenas, but little is known about whether courtship proficiency changes with experience and/or whether subordinates actively contribute to enhancing the resident male's mating success. In this study, we investigated male-male associations in wild spotted bowerbirds (Ptilonorhynchus maculatus). We first sought to determine whether courtship behaviour differs based on bower ownership status. We then examined whether social interactions between bower owners and subordinate males may qualify as courtship coalitions. Our analysis of courtship postural components did not reveal differences in timing or relative occurrence of postural components between subordinate males and bower owners, whereas we found evidence that male-male associations in spotted bowerbirds may provide an example of rudimentary courtship coalitions. In particular, higher subordinate attendance is associated with lower destruction rates by neighbouring rivals and with overall higher mating success, and male pairs are stable in subsequent years. This study provides novel information about social dynamics among male bowerbirds, and further insights into the evolution of coalitionary behaviour in male displays. Significance statement: Same-sex associations between established males and subordinate visitors on display arenas are common in birds, yet poorly understood. Using video recordings from a population of wild spotted bowerbirds, Ptilonorhynchus maculatus, we performed a quantitative analysis on motor courtship components across males, and on their social interactions on display arenas to investigate the nature of male-male partnerships. Our results showed that motor courtship performance in subordinate visitors is not suggestive of an early ontogenetic stage, as previously speculated. Moreover, though bower 'owners' and subordinate males do not coordinate their behaviour during courtship or bower building, male-male partnerships may qualify as a rudimentary or incipient form of courtship coalitions. Subordinate males are tolerated at bowers, the magnitude of subordinate attendance correlates with owner males' mating success, and repeated interactions between individuals reveal consistent partner associations. Supplementary Information: The online version contains supplementary material available at 10.1007/s00265-022-03200-x.

epg5 knockout leads to the impairment of reproductive success and courtship behaviour in a zebrafish model of autophagy-related diseases.

BACKGROUND: Dysregulation of the autophagic flux is linked to a wide array of human diseases, and recent findings highlighted the central role of autophagy in reproduction, as well as an association between impairment of autophagy and behavioural disorders. Here we deepened on the possible multilevel link between impairment of the autophagic processes and reproduction at both the physiological and the behavioural level in a zebrafish mutant model. METHODS: Using a KO epg5 zebrafish line we analysed male breeding success, fertility rate, offspring survival, ejaculate quality, sperm and testes morphology, and courtship behaviour. To this aim physiological, histological, ultrastructural and behavioural analyses on epg5+/+ and mutant epg5-/- males coupled to WT females were applied. RESULTS: We observed an impairment of male reproductive performance in mutant epg5-/- males that showed a lower breeding success with a reduced mean number of eggs spawned by their WT female partners. The spermatogenesis and the ability to produce fertilising ejaculates were not drastically impaired in our mutant males, whereas we observed a reduction of their courtship behaviour that might contribute to explain their lower overall reproductive success. CONCLUSION: Collectively our findings corroborate the hypothesis of a multilevel link between the autophagic process and reproduction. Moreover, by giving a first glimpse on behavioural disorders associated to epg5 KO in model zebrafish, our results open the way to more extensive behavioural analyses, also beyond the reproductive events, that might serve as new tools for the molecular screening of autophagy-related multisystemic and neurodegenerative diseases.

Reproductive roles of the vasopressin/oxytocin neuropeptide family in teleost fishes.

The vertebrate nonapeptide families arginine vasopressin (AVP) and oxytocin (OXT) are considered to have evolved from a single vasopressin-like peptide present in invertebrates and termed arginine vasotocin in early vertebrate evolution. Unprecedented genome sequence availability has more recently allowed new insight into the evolution of nonapeptides and especially their receptor families in the context of whole genome duplications. In bony fish, nonapeptide homologues of AVP termed arginine vasotocin (Avp) and an OXT family peptide (Oxt) originally termed isotocin have been characterized. While reproductive roles of both nonapeptide families have historically been studied in several vertebrates, their roles in teleost reproduction remain much less understood. Taking advantage of novel genome resources and associated technological advances such as genetic modifications in fish models, we here critically review the current state of knowledge regarding the roles of nonapeptide systems in teleost reproduction. We further discuss sources of plasticity of the conserved nonapeptide systems in the context of diverse reproductive phenotypes observed in teleost fishes. Given the dual roles of preoptic area (POA) synthesized Avp and Oxt as neuromodulators and endocrine/paracrine factors, we focus on known roles of both peptides on reproductive behaviour and the regulation of the hypothalamic-pituitary-gonadal axis. Emphasis is placed on the identification of a gonadal nonapeptide system that plays critical roles in both steroidogenesis and gamete maturation. We conclude by highlighting key research gaps including a call for translational studies linking new mechanistic understanding of nonapeptide regulated physiology in the context of aquaculture, conservation biology and ecotoxicology.

Reproductive potential of mosquitofish is reduced by the masculinizing effect of a synthetic progesterone, gestodene: Evidence from morphology, courtship behaviour, ovary histology, sex hormones and gene expressions.

The ever-increasing use of synthetic hormones, especially progestins, for medical applications has drawn growing concerns due to their potential endocrine disrupting effects that may diminish the reproductive outputs of aquatic organisms. Using mosquitofish (Gambusia affinis) as a model species, we tested whether gestodene (GES), a commonly used progestin, can alter the expressions of genes associated with sex hormone synthesis and cause ensuing changes in morphological features, courtship behaviour and oocyte development. After exposing to GES at environmentally relevant concentrations (2.96, 32.9 and 354 ng L-1) for 40 days, we found that GES, especially at 354 ng L-1, induced masculinization of female fish, indicated by the reduced body weight to length ratio and development of gonopodia (i.e. anal fins of male fish). Thus, the males showed less intimacy and mating interest towards the GES-exposed females, indicated by the reduced time spent on attending, following and mating behaviours. While oocyte development was seemingly unaffected by GES, spermatogonia were developed in the ovary. All the aforementioned masculinizing effects of GES were associated with the increased testosterone level and decreased estradiol level, driven by upregulating androgen receptor genes (Arα and Arß). Overall, our findings suggest that progestins could undermine the reproductive potential of aquatic organisms and hence their persistence in the progestin-contaminated environment.

Low frequencies in the display vocalization of the Western Capercaillie (Tetrao urogallus).

Only a few bird species are known to produce low-frequency vocalizations. We analyzed the display vocalizations of Western Capercaillie males kept in breeding centers and identified harmonically structured signals with a fundamental frequency of 28.7 ± 1.2 Hz (25.6-31.6 Hz). These low-frequency components temporally overlap with the Whetting phase (96% of its duration) and they significantly contribute to the distinct vocal expression between individuals. The resulting model of discrimination analysis classified 67.6% vocalizations (63%, cross-validated result) correctly to the specific individual in comparison to the probability by chance of 12.5%. We discuss a possible function of low-frequency components that remains unclear. The occurrence of such low frequencies is surprising as this grouse is substantially smaller than cassowaries (Southern cassowary Casuarius casuarius and Dwarf cassowary Casuarius bennetti) , the species that produces similarly low frequencies. Because these low frequency components temporarily overlap with the Whetting phase, they are hardly audible from a distance larger than several meters.

A reproductive role for the nonapeptides vasotocin and isotocin in male zebrafish (Danio rerio).

Two distinct nonapeptide systems, vasotocin- and oxytocin-related peptides, evolved in vertebrates. Their role in male zebrafish reproduction has not been formally investigated. We hypothesized that the teleost nonapeptides vasotocin and isotocin stimulate male zebrafish reproductive physiology and success by affecting central neuronal and/or peripheral endocrine pathways. Pharmacological inhibition experiments revealed that both vasotocin and isotocin contribute significantly to male reproductive success, which in the case of vasotocin correlated significantly with indices of male courtship behavior. Interestingly, co-administration of vasotocin and isotocin antagonists completely abolished male reproductive success without affecting male courtship behavior and endocrine indices, possibly linked to a synergistic action of nonapeptides on male pheromone release. To further probe the nonapeptides' role in male zebrafish reproduction, we subsequently tested whether male zebrafish nonapeptide systems were acutely activated by the female releaser pheromone PGF2α, a strong chemoattractant and important reproductive cue in males which stimulates courtship behavior. Male zebrafish attracted to PGF2α in a choice assay exhibited acute increases in neuronal activation marker p-ERK immunoreactivity in the ventral glomerulus of the olfactory bulb and the preoptic area, however no co-localization with isotocin was observed. Conversely, PGF2α time-dependently stimulated whole brain isotocin mRNA abundance, suggesting secondary longer-term effects of PGF2α exposure on the central isotocinergic system. While the current lack of vasotocin-specific antibodies for zebrafish does not allow to probe acute activation of vasotocinergic neurons, whole brain vasotocin mRNA was not significantly affected by PGF2α exposure. Together, our results identify a role for nonapeptides in male zebrafish reproductive physiology and success.

Signalling interactions during ontogeny are a cause of social plasticity in Enchenopa treehoppers (Hemiptera: Membracidae).

We recently discovered that there is a social ontogeny of signals and preferences in Enchenopa treehoppers. Nymphs signalled throughout their development; some signal features changed gradually and in sexually dimorphic ways throughout ontogeny; and some adult male signal features and female mate preferences differed between individuals reared in isolation or groups. In this paper, we investigate whether signalling interactions during ontogeny are a cause of plasticity in mating signals and preferences. We subjected Enchenopa nymphs to treatments of either: rearing in aggregations (the natural condition), in isolation, or in isolation with playbacks of nymph signals. We then described variation in the signals and mating preferences of individuals that developed in those conditions. The playback treatments partially "rescued" the signal and preference phenotypes, resulting in phenotypes either similar to those that result from rearing in aggregations, or intermediate between those that result from rearing in isolation or in aggregations. These results pin-point signalling interactions during ontogeny as an important cause of plasticity in signals and mate preferences.

Male prairie voles with different avpr1a microsatellite lengths do not differ in courtship behaviour.

Females are generally expected to be selective when choosing their social and sexual partners. In a previous laboratory study, female prairie voles (Microtus ochrogaster) showed significant social and sexual preferences for males with longer microsatellite DNA within the avpr1a gene encoding the vasopressin 1a receptor, as predicted if females select mates whose parental behaviour should increase female reproductive success. We tested the hypothesis that males with short versus long avpr1a microsatellite alleles exhibit differences in courtship behaviour, which could act as cues for female mate preference. The only behavioural difference we detected between males with short versus long avpr1a microsatellite alleles in mate preference trials was that males with short avpr1a microsatellite alleles sniffed the anogenital region of females more frequently during the first two days of the trials. Our results did not strongly support the hypothesis that a male's avpr1a genotype predicts the courtship behaviours we measured and suggests that other courtship behaviours or traits, such as odour and vocalizations, may be more important to female prairie voles when choosing mates. Additional studies using a wider array of species are needed to assess the degree to which male mammal courtship behaviour provides information on mate quality to females.

On the function of the ultimate legs of some Scolopendridae (Chilopoda, Scolopendromorpha).

The function of the variously shaped ultimate legs of Scolopendridae is briefly reviewed. Their function in Scolopendraheros Girard, 1853, Scolopendrasubspinipes Leach, 1815, Scolopendramorsitans (Linnaeus, 1758), Scolopendragalapagoensis Bollman, 1889, Scolopendrahainanum Kronmüller, 2012, Scolopendraspinosissima Kraepelin, 1903 Cormocephalusaurantiipes (Newport, 1844) and Ethmostigmustrigonopodus (Leach, 1817), in which they are least specialised has been investigated. Specimens were tapped with forceps on different parts of the trunk to simulate the attack of a predator. When tapped on the first third of the trunk (near the head), the centipedes attacked the forceps with their forcipules. When tapped on the last third or the ultimate legs, they adopted a warning position, raising the ultimate legs to display the ventral and medial prefemoral spines as well as the spined coxopleural processes. In some cases the centipedes attacked the forceps with the claws of the ultimate legs by chopping down on them after lifting the legs high into the warning position. When tapped in the mid part of the trunk, the centipedes curled sideways to reach the forceps with their forcipules and ultimate legs simultaneously. Scolopendragalapagoensis not only lifted the ultimate legs into the warning position but also the last 3-4 pairs of locomotory legs, presenting their distodorsal prefemoral spines. This resembles the warning posture of some spiders. In addition to their function in warning behaviour, defensive stabbing, ritualised meeting reactions and during courtship behaviour, the ultimate legs may in addition act as hooks and perhaps be involved in species recognition. No evidence was found that the ultimate legs are used to catch prey, nor of prey or predators being held between the prefemora.

Review: Improving our knowledge of male mosquito biology in relation to genetic control programmes.

The enormous burden placed on populations worldwide by mosquito-borne diseases, most notably malaria and dengue, is currently being tackled by the use of insecticides sprayed in residences or applied to bednets, and in the case of dengue vectors through reduction of larval breeding sites or larviciding with insecticides thereof. However, these methods are under threat from, amongst other issues, the development of insecticide resistance and the practical difficulty of maintaining long-term community-wide efforts. The sterile insect technique (SIT), whose success hinges on having a good understanding of the biology and behaviour of the male mosquito, is an additional weapon in the limited arsenal against mosquito vectors. The successful production and release of sterile males, which is the mechanism of population suppression by SIT, relies on the release of mass-reared sterile males able to confer sterility in the target population by mating with wild females. A five year Joint FAO/IAEA Coordinated Research Project brought together researchers from around the world to investigate the pre-mating conditions of male mosquitoes (physiology and behaviour, resource acquisition and allocation, and dispersal), the mosquito mating systems and the contribution of molecular or chemical approaches to the understanding of male mosquito mating behaviour. A summary of the existing knowledge and the main novel findings of this group is reviewed here, and further presented in the reviews and research articles that form this Acta Tropica special issue.