The spitting behavior of two species of spitting cobras

Spitting cobras defend themselves by spitting their venom in the face of a harasser. Although it is common belief that spitting cobras direct their venom at the eyes of an aggressor, this has never been investigated. Here, we show that the spitting act of cobras (Naja nigricollis and N. pallida) can readily be triggered by a moving human face or by a moving real size photo of a human face. In contrast, a stationary human face (real or photo) or a moving or stationary human hand does not trigger the spitting act. If threatened, spitting cobras aim their venom, ejected either in two distinct jets (N. pallida) or in a fine spray (N. nigricollis), either between the eyes or at one eye. In both cobra species investigated, the width and height of the area hit by the venom was independent of eye distance (test range 5.5 cm and 11 cm). During the spitting act the cobras performed fast undulating head movements that lead to a larger distribution of their venom. This behavior increases the probability that at least one eye of the aggressor is hit. Electronic Supplementary Material Supplementary material is available for this article at     

Spitting cobras adjust their venom distribution to target distance

If threatened by a human, spitting cobras defend themselves by ejecting their venom toward the face of the antagonist. Circulating head movements of the cobra ensure that the venom is distributed over the face. To assure an optimal distribution of the venom, the amplitudes of head movements should decrease with increasing target distance. To find out whether cobras (Naja pallida and N. nigricollis) adjust their spitting behavior according to target distance we induced spitting from different distances and analyzed their spitting patterns. Our results show that the spray pattern of spiting cobras is not fixed. Instead the snake matches its venom distribution to the size of the target independent of target distance.     

The phylogeny of cobras inferred from mitochondrial DNA sequences: evolution of venom spitting and the phylogeography of the African spitting cobras (Serpentes: Elapidae: Naja nigricollis complex)

We use phylogenetic analysis of 1333 bp of mitochondrial DNA sequence to investigate the phylogeny and historical biogeography of the cobra-like elapid snakes, with special reference to the evolution of spitting and the phylogeography of the African spitting cobras, a radiation widespread in open vegetational formations throughout sub-Saharan Africa. Our results suggest that spitting adaptations appear to have evolved three times in cobras, but alternative scenarios cannot be rejected. The Asiatic Naja are monophyletic and originate from a single colonization of Asia from Africa. The radiation of the African spitting Naja appears to date back to the early Miocene and many speciation events in the group predate the Pliocene expansion of grasslands and the radiation of large grazing mammals in Africa. The cladogenic events in this complex appear to have been triggered by both ecological changes and tectonic events associated with the formation and expansion of the African Rift Valley. Taxonomically, our data confirm the inclusion of Boulengerina and Paranaja within Naja, and reveal a clade of African rainforest cobras including N. melanoleuca, Paranaja multifasciata and Boulengerina that constitutes the sister clade of the African open-formation non-spitting cobras. Naja nigricollis is polyphyletic, and we therefore recognize N. nigricincta as a separate species, more closely related to N. ashei and N. mossambica than to N. nigricollis.     

A spitting image: specializations in archerfish eyes for vision at the interface between air and water

Archerfish are famous for spitting jets of water to capture terrestrial insects, a task that not only requires oral dexterity, but also the ability to detect small camouflaged prey against a visually complex background of overhanging foliage. Because detection of olfactory, auditory and tactile cues is diminished at air–water interfaces, archerfish must depend almost entirely on visual cues to mediate their sensory interactions with the aerial world. During spitting, their eyes remain below the water''s surface and must adapt to the optical demands of both aquatic and aerial fields of view. These challenges suggest that archerfish eyes may be specially adapted to life at the interface between air and water. Using microspectrophotometry to characterize the spectral absorbance of photoreceptors, we find that archerfish have differentially tuned their rods and cones across their retina, correlated with spectral differences in aquatic and aerial fields of view. Spatial resolving power also differs for aquatic and aerial fields of view with maximum visual resolution (6.9 cycles per degree) aligned with their preferred spitting angle. These measurements provide insight into the functional significance of intraretinal variability in archerfish and infer intraretinal variability may be expected among surface fishes or vertebrates where different fields of view vary markedly.     

The ghost of a recent invasion in the reduced feeding rates of spitting cobras during the dry season in a rainforest region of tropical Africa?

Two species of cobras (Naja melanoleuca and Naja nigricollis) are known to occur in south eastern Nigeria, where much of the pristine rainforest surface has been felled in the last thirty years, and where the actual landscape is generally constituted by a mosaic of farmlands, plantations, suburban areas, with a few remnant forest fragments. In this region, Naja nigricollis is currently extending its range, especially by exploiting recently deforested areas. Based on the known general distribution range of this species and on the available literature data, it appears that Naja nigricollis has been colonizing the forested region of south eastern Nigeria, starting from the relatively arid savannas of central Nigeria, where this species aestivates during the driest months. In the forest region, however, snakes do not need to aestivate during the dry season. Nevertheless, whereas Naja melanoleuca has a foraging activity extended all-the-year round, Naja nigricollis reduces feeding rates during the dry months, although it does not suspend above-ground activity in these months. I suggest that rainforest spitting cobras suspend feeding during the dry months because their behaviour is just a ‘ghost’ of their recent past, when they were ‘normal’ spitting cobras of dry savana regions, which were thus constrained to aestivate during the dry season as it is the rule in this species in central and northern Nigeria. The ‘gost-of-the-past hypothesis’ seems to fit well with the ‘invading’ presence of Naja nigricollis in Nigerian areas where they were reported as rare or, even, absent, up to a few decades ago. Other hypotheses are discussed, and rejected.     

Stimulus control of predatory behaviour in red spitting cobras (Naja mossambica pallida) and prairie rattlesnakes (Crotalus v. viridis)

Two experiments examined the responses of red spitting cobras (Naja mossambica pallida) to stimulus properties of rodent prey. In experiment 1 cobras exhibited elevated rates of tongue flicking (RTF) and trail-following behaviour after striking mice and after 15-s presentations of visual-chemical-thermal cues arising from mice held just out of striking range. Disturbance with a clean snake hook did not produce similar effects. In experiment 2 these cobras were compared with prairie rattlesnakes (Crotalus v. viridis). All snakes were exposed to five stimulus conditions in their home cages: (1) 3 s of disturbance (D) with a clean snake hook; (2) 15 s of D; (3) 3 s inspection of a live mouse held out of striking range (NS); (4) 15 s of NS; and (5) a presentation of a live mouse inside striking range (S). The snakes always struck in the latter condition, usually within 3 s. Rattlesnakes exhibited elevated RTF only after S, whereas cobras did so after 15 s of both NS and S. The performance of rattlesnakes was consistent with the assumption that these animals are ambushers; that of cobras was consistent with the assumption that these animals are active hunters which will chase and catch visually detected prey.     

Pentastomes (Pentastomida, Armillifer armillatus Wyman, 1848) in snakes from Zambia

Twenty-three snakes, belonging to eight different species, were collected from rural areas of Zambia and inspected for the presence of pentastomes. Pentastomid parasites were found in four snakes: one African rock python (Python sebae), one puff adder (Bitis arietans) and two Mozambique spitting cobras (Naja mossambica) were infested with a small number of Armillifer armillatus, respectively five, two and one adult parasites. As humans can be incidental/intermediate hosts for reptilian pentastomes, the zoonotic potential of these parasites, especially in tropical countries, is discussed.     

Does the perception of moving eyes trigger reflexive visual orienting in autism?

Does movement of the eyes in one or another direction function as an automatic attentional cue to a location of interest? Two experiments explored the directional movement of the eyes in a full face for speed of detection of an aftercoming location target in young people with autism and in control participants. Our aim was to investigate whether a low-level perceptual impairment underlies the delay in gaze following characteristic of autism. The participants'' task was to detect a target appearing on the left or right of the screen either 100 ms or 800 ms after a face cue appeared with eyes averting to the left or right. Despite instructions to ignore eye-movement in the face cue, people with autism and control adolescents were quicker to detect targets that had been preceded by an eye movement cue congruent with target location compared with targets preceded by an incongruent eye movement cue. The attention shifts are thought to be reflexive because the cue was to be ignored, and because the effect was found even when cue-target duration was short (100 ms). Because (experiment two) the effect persisted even when the face was inverted, it would seem that the direction of movement of eyes can provide a powerful (involuntary) cue to a location.     

Baby Schema in Infant Faces Induces Cuteness Perception and Motivation for Caretaking in Adults

Ethologist Konrad Lorenz proposed that baby schema ('Kindchenschema') is a set of infantile physical features such as the large head, round face and big eyes that is perceived as cute and motivates caretaking behavior in other individuals, with the evolutionary function of enhancing offspring survival. Previous work on this fundamental concept was restricted to schematic baby representations or correlative approaches. Here, we experimentally tested the effects of baby schema on the perception of cuteness and the motivation for caretaking using photographs of infant faces. Employing quantitative techniques, we parametrically manipulated the baby schema content to produce infant faces with high (e.g. round face and high forehead), and low (e. g. narrow face and low forehead) baby schema features that retained all the characteristics of a photographic portrait. Undergraduate students (n = 122) rated these infants' cuteness and their motivation to take care of them. The high baby schema infants were rated as more cute and elicited stronger motivation for caretaking than the unmanipulated and the low baby schema infants. This is the first experimental proof of the baby schema effects in actual infant faces. Our findings indicate that the baby schema response is a critical function of human social cognition that may be the basis of caregiving and have implications for infant–caretaker interactions.     

Cover-seeking behavior in red spitting cobras (Naja mossambica pallida): Effects of tactile cues and darkness

Eleven red spitting cobras, Naja mossambica pallida, used clear Plexiglas hiding boxes as frequently as they used dark ones in a successive discrimination paradigm (experiment 1), which indicated that thigmotaxic cues can satisfy the cover-seeking needs of the snakes. In simultaneous discrimination tests, however, dark places were always preferred by the snakes (experiments II, III). Therefore, although thigmotaxic cues are sufficient, these plus darkness constitute a more favorable alternative for N m pallida. The husbandry advantages associated with clear hiding boxes, together with their sufficiency for the snakes, argue strongly for their use in many captive environments.     

"X-ray vision" and the evolution of forward-facing eyes

Why do our eyes face forward, and why do many mammals have eyes facing sideways? Here, we describe results suggesting that the degree of binocular convergence is selected to maximize how much the mammal can see in its environment. Mammals in non-cluttered environments can see the most around them with panoramic, laterally directed eyes. Mammals in cluttered environments, however, can see best when their eyes face forward, for binocularity has the power of "seeing through" clutter out in the world. Evidence across mammals closely fits the predictions of this "X-ray" hypothesis.     

Attentiveness to eyes predicts generosity in a reputation-relevant context

Eyes play a vital role in human social interactions. In fact, some prior work indicates that simply the presence of eyes or eye-like stimuli increases people's prosocial behavior, arguably because the eyes serve as cues of being watched and thus elicit reputational concerns. The current study was designed to address two questions in this regard. First, we examined how salient the eyes are among the human features. Second, we asked whether individual differences in attentiveness to eyes (but not other human features such as ears or hands) are predictive of reputation-enhancing behavior. Using an eye-tracking paradigm, we found that participants looked longest to eyes compared to other human features. Critically, greater attentiveness to eyes correlated with greater generosity on a donation task, but only in a reputation-relevant context (i.e., when donations were public but not when they were anonymous). Attentiveness to other human features did not predict donation behavior. Eyes are thus an especially salient human feature, and attentiveness to eyes may signal individuals' concerns about their reputations.     

Exceptional Variation on a Common Theme: The Evolution of Crustacean Compound Eyes

The Crustacea contain an amazing, and often (to humans) bizarre, array of visual designs. This diversity includes many different examples of both simple and compound eyes, each with standard or uniquely crustacean features. In this review, we focus on the anatomical variation, optical principles, and molecular diversity of crustacean compound eyes to illustrate how the complicated structures involved in vision are adapted for particular environments. Using this knowledge as a starting point, and considering what is known of crustacean evolution overall, we present the most recent ideas of how crustacean compound eyes have evolved and show how eyes that are based on fundamentally different optical principles can in fact be derived from each other and thus be closely related through common descent.     

Shyness in Early Infancy: Approach-Avoidance Conflicts in Temperament and Hypersensitivity to Eyes during Initial Gazes to Faces

‘Infant shyness’, in which infants react shyly to adult strangers, presents during the third quarter of the first year. Researchers claim that shy children over the age of three years are experiencing approach-avoidance conflicts. Counter-intuitively, shy children do not avoid the eyes when scanning faces; rather, they spend more time looking at the eye region than non-shy children do. It is currently unknown whether young infants show this conflicted shyness and its corresponding characteristic pattern of face scanning. Here, using infant behavioral questionnaires and an eye-tracking system, we found that highly shy infants had high scores for both approach and fear temperaments (i.e., approach-avoidance conflict) and that they showed longer dwell times in the eye regions than less shy infants during their initial fixations to facial stimuli. This initial hypersensitivity to the eyes was independent of whether the viewed faces were of their mothers or strangers. Moreover, highly shy infants preferred strangers with an averted gaze and face to strangers with a directed gaze and face. This initial scanning of the eye region and the overall preference for averted gaze faces were not explained solely by the infants’ age or temperament (i.e., approach or fear). We suggest that infant shyness involves a conflict in temperament between the desire to approach and the fear of strangers, and this conflict is the psychological mechanism underlying infants’ characteristic behavior in face scanning.     

舟山眼镜蛇对光暗周期加热光源反应所导致的体温变化

用 7条舟山眼镜蛇 (Najaatra)研究动物对光暗周期加热光源反应所导致的体温变化。设计两项实验 ,每项实验历时 9d。实验一光照期覆盖整个白天 ,实验二光照期覆盖整个晚上。眼镜蛇仅在加热光源开启期间进行体温调节 ,但光照期内任何阶段都未发现所有个体同时处于热活动状态。两项实验中热活动个体百分比的时间变化显著 ,实验二热活动个体百分比波动相对大于实验一。在加热光源开启期间 ,实验一热活动眼镜蛇的百分比总体上大于实验二。在两项实验中 ,热活动眼镜蛇体温的时间变化都不显著。实验一热活动眼镜蛇的体温高于实验二 ,而两项实验中不处于热活动状态的眼镜蛇的平均体温无显著差异。实验一热活动眼镜蛇 (31 1±0 8°C)选择的体温上限高于实验二眼镜蛇 (2 6 0± 0 9°C)。在两项实验的任何时间段内 ,眼镜蛇的体温都不低于环境温度。     

Attractiveness qualifies the effect of observation on trusting behavior in an economic game

Recent studies show that subtle cues of observation affect cooperation even when anonymity is explicitly assured. For instance, recent studies have shown that the presence of eyes increases cooperation on social economic tasks. Here, we tested the effects of cues of observation on trusting behavior in a two-player Trust game and the extent to which these effects are qualified by participants' own attractiveness. Although explicit cues of being observed (i.e., when participants were informed that the other player would see their face) tended to increase trusting behavior, this effect was qualified by the participants' other-rated attractiveness (estimated from third-party ratings of face photographs). Participants' own physical attractiveness was positively correlated with the extent to which they trusted others more when they believed they could be seen than when they believed they could not be seen. This interaction between cues of observation and own attractiveness suggests context dependence of trusting behavior that is sensitive to whether and how others react to one's physical appearance.     

The use of different eye regions in the mantis shrimp Hemisquilla californiensis Stephenson, 1967 (Crustacea: Stomatopoda) for detecting objects

A behavioral assay was used to assess the ability of the stomatopod Hemisquilla californiensis to perceive and respond to a moving target under different wavelengths and intensities of light illumination. Subjects responded to targets rotating horizontally across their visual field by a brief startle response of their eyes or antennules but did not track the targets. Under white light responses were elicited down to a light intensity of 0.9 μW cm^− 2. Responses were seen in blue light at intensities as low as 0.5 μW cm^− 2, and in green light down to 1.0 μW cm^− 2. The animals were less sensitive to red light, with no responses seen at intensities below 3.0 μW cm^− 2. Subjects did not respond to the targets at all under infrared light. This response pattern mirrors the computed sensitivity spectrum of ommatidia in the species' peripheral hemispheres but not that in most of the central bands. We conclude that this species uses the monochromatic vision in the peripheral hemispheres of its eyes to recognize objects and that the sharply tuned color receptors of the central band serve to add supplemental information if light conditions allow.     

One eye but no vision: cave fish with induced eyes do not respond to light

One of the most intriguing questions in evolutionary biology is the degree to which behavior is a necessary consequence of morphology. We explore this issue by examining phototactic behavior in epigean (eyed surface-dwelling) and troglomorphic (blind cave) forms of the teleost Astyanax fasciatus whose eyes were modified during embryogenesis by removing one or both lens vesicles from the epigean form or by transplanting the lens vesicle from an epigean fish into the optic cup of a blind cave form. Lens removal results in eye degeneration and blindness in adult epigean fish, whereas lens transplantation stimulates growth of the eye, inducing the development of optic tissues in the normally eyeless adult cave fish. Photoresponsiveness was examined by placing fish in an aquarium with one half illuminated and the other half dark and scoring their presence in the illuminated or dark half. Both the eyeless epigean fish and cave fish with induced eyes are indifferent to the illumination whereas the surface forms are scotophilic, suggesting that optic development and phototactic behavior are decoupled.     

The Role of Eyespots in Predator Recognition and Antipredatory Behaviour of the Paradise Fish,Macropodus opercularis L.

The behaviour of the paradise fish in the presence of either a living pike or catfish or various dummies or a catfish equipped with a pair of artificial big eyes was studied. Analyzing the first encounter two separate periods of the predator recognition process could be identified. The primary form of the antipredatory reaction was orientation (looking at the object while keeping a constant eye and body position) which was elicited by any fish-like bodies but not by bare eyes without a body. During orientation the presence of pike or catfish equipped with artificial eyes or a dummy with horizontal eyes elicited fin erecting display. This reaction was sporadic in the presence of a normal catfish which has very small eyes. The biological significance of this separate organization of exploration is discussed.     

Heterologous expression,characterization and evaluation of the matrix protein from Newcastle disease virus as a target for antiviral therapies

Newcastle disease virus (NDV) is an infectious agent of a large variety of birds, including chicken, which poses a real threat to the agriculture industry. Matrix (M) proteins of NDV and many other viruses perform critical functions during viral assembly and budding from the host cell. M-proteins are well conserved and therefore are potential targets for antiviral therapies. To validate this, we expressed the NDV M-protein in its native form in Saccharomyces cerevisiae and in inclusion bodies in Escherichia coli. Proper refolding of the recombinant protein produced in E. coli was verified using circular dichroism and infrared spectroscopies and electron microscopy. Immunization of chickens with the NDV M-protein elicited significant serum antibody titers. However, the antibodies conferred little protection against the ND following lethal viral challenges. We conclude that the M-protein is not exposed on the surface of the host cell or the virus at any stage during its life cycle. We discuss how the conserved M-protein can further be exploited as an antiviral drug target.