Morphology and spectral sensitivity of long visual fibers and lamina monopolar cells in the butterfly Papilio xuthus.

Extensive analysis of the flower-visiting behavior of a butterfly, Papilio xuthus, has indicated complex interaction between chromatic, achromatic, and motion cues. Their eyes are spectrally rich with six classes of photoreceptors, respectively sensitive in the ultraviolet, violet, blue, green, red, and broad-band wavelength regions. Here, we studied the anatomy and physiology of photoreceptors and second-order neurons of P. xuthus, focusing on their spectral sensitivities and projection terminals to address where the early visual integration takes place. We thus found the ultraviolet, violet, and blue photoreceptors and all second-order neurons terminate in the distal region of the second optic ganglion, the medulla. We identified five types of second-order neurons based on the arborization in the first optic ganglion, the lamina, and the shape of the medulla terminals. Their spectral sensitivity is independent of the morphological types but reflects the combination of pre-synaptic photoreceptors. The results indicate that the distal medulla is the most plausible region for early visual integration.

Elongating, entwining, and dragging: mechanism for adaptive locomotion of tubificine worm blobs in a confined environment.

Worms often aggregate through physical connections and exhibit remarkable functions such as efficient migration, survival under environmental changes, and defense against predators. In particular, entangled blobs demonstrate versatile behaviors for their survival; they form spherical blobs and migrate collectively by flexibly changing their shape in response to the environment. In contrast to previous studies on the collective behavior of worm blobs that focused on locomotion in a flat environment, we investigated the mechanisms underlying their adaptive motion in confined environments, focusing on tubificine worm collectives. We first performed several behavioral experiments to observe the aggregation process, collective response to aversive stimuli, the motion of a few worms, and blob motion in confined spaces with and without pegs. We found the blob deformed and passed through a narrow passage using environmental heterogeneities. Based on these behavioral findings, we constructed a simple two-dimensional agent-based model wherein the flexible body of a worm was described as a cross-shaped agent that could deform, rotate, and translate. The simulations demonstrated that the behavioral findings were well-reproduced. Our findings aid in understanding how physical interactions contribute to generating adaptive collective behaviors in real-world environments as well as in designing novel swarm robotic systems consisting of soft agents.

Connectome of the lamina reveals the circuit for early color processing in the visual pathway of a butterfly.

Connectomics has become a standard neuroscience methodology in a few model animals,1 with the visual system being a popular target of study.2-5 Combining connectomics with circuit and behavioral physiology, recent studies on the color vision of the fruit fly Drosophila melanogaster have focused on the mechanisms underlying early wavelength processing in the optic ganglia.6-8 However, the color vision capabilities of D. melanogaster are limited,9 compared with many flower-visiting insects.10,11 For example, a butterfly Papilio xuthus has six spectral classes of photoreceptors. Each ommatidium contains nine photoreceptors in one of three fixed combinations, making the eye an array of three spectrally distinct ommatidia types.12 Behaviorally, P. xuthus can detect 1 nm differences in light wavelength across the spectrum from ultraviolet to red, outperforming humans.13 What is the neuronal basis of such precise color vision? How does such a system evolve? Addressing these questions requires comparative studies at the circuit level. Here, we performed a connectome analysis in the first optic ganglion, the lamina, of P. xuthus. The lamina comprises cartridges, each typically containing nine photoreceptor axons from a single ommatidium and four second-order neurons. We found abundant inter-photoreceptor connections, which are absent in the lamina of D. melanogaster. We also identified connections between neighboring cartridges, particularly those receiving inputs from spectrally distinct ommatidia. The linear summation of synaptic connections well explains the spectral sensitivity of photoreceptors and second-order neurons in the lamina.

A general model of locomotion of brittle stars with a variable number of arms.

Typical brittle stars have five radially symmetrical arms that coordinate to move the body in a certain direction. However, some species have a variable number of arms, which is a unique trait since intact animals normally have a fixed number of limbs. How does a single species manage different numbers of appendages for adaptive locomotion? We herein describe locomotion in Ophiactis brachyaspis with four, five, six and seven arms to propose a common rule for the movement of brittle stars with different numbers of arms. For this, we mechanically stimulated one arm of individuals to analyse escape direction and arm movement. By gathering quantitative indices and employing Bayesian statistical modelling, we noted a pattern: regardless of the total number of arms, an anterior position emerges at one of the second neighbouring arms to a mechanically stimulated arm, while arms adjacent to the anterior one synchronously work as left and right rowers. We propose a model in which an afferent signal runs clockwise or anticlockwise along the nerve ring while linearly counting how many arms it passes through. With this model, the question on how 'left and right' emerges in a radially symmetrical body via a decentralized system is answered.

Sea stars of the genus Henricia Gray, 1840 (Echinodermata, Asteroidea) from Vostok Bay, Sea of Japan.

We report seven species of the genus Henricia Gray, 1840 that were found in Vostok Bay, and two species from adjacent area, known from museum collection or seen in underwater footage. while existing literature reported no confirmed species from this area. Most of these species: H. djakonovi, H. alexeyi, H. densispina, H. hayashii, H. granulifera, H. pacifica, H. asiatica, and H. oculata robusta were reported from the Sea of Japan previously. H. nipponica, known from Japan, is reported from Russian seas for the first time. All studied taxa are re-described here using a range of morphological characters and partial 16S rRNA nucleotide sequences, life colorations of several species are reported for the first time, and an identification key is provided. Lectotype designations are fixed for studied series of species described by AM Djakonov.

Different Synchrony in Rhythmic Movement Caused by Morphological Difference between Five- and Six-armed Brittle Stars.

Physiological experiments and mathematical models have supported that neuronal activity is crucial for coordinating rhythmic movements in animals. On the other hand, robotics studies have suggested the importance of physical properties made by body structure, i.e. morphology. However, it remains unclear how morphology affects movement coordination in animals, independent of neuronal activity. To begin to understand this issue, our study reports a rhythmic movement in the green brittle star Ophiarachna incrassata. We found this animal moved five radially symmetric parts in a well-ordered unsynchronized pattern. We built a phenomenological model where internal fluid flows between the five body parts to explain the coordinated pattern without considering neuronal activity. Changing the number of the body parts from five to six, we simulated a synchronized pattern, which was demonstrated also by an individual with six symmetric parts. Our model suggests a different number in morphology makes a different fluid flow, leading to a different synchronization pattern in the animal.