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Filter-feeding has been present for hundreds of millions of years, independently evolving in aquatic vertebrates' numerous times. Mysticete whales are a group of gigantic, marine filter-feeders that are defined by their fringed baleen and are divided into two groups: balaenids and rorquals. Recent studies have shown that balaenids likely feed using a self-cleaning, cross-flow filtration mechanism where food particles are collected and then swept to the esophagus for swallowing. However, it is unclear how filtering is achieved in the rorquals (Balaenopteridae). Lunging rorqual whales engulf enormous masses of both prey and water; the prey is then separated from the water through baleen plates lining the length of their upper jaw and positioned perpendicular to flow. Rorqual baleen is composed of both major (larger) and minor (smaller) keratin plates containing embedded fringe that extends into the whale's mouth, forming a filtering fringe. We used a multimodal approach, including microcomputed tomography (µCT) and scanning electron microscopy (SEM), to visualize and describe the variability in baleen anatomy across five species of rorqual whales, spanning two orders of magnitude in body length. For most morphological measurements, larger whales exhibited hypoallometry relative to body length. µCT and SEM revealed that the major and minor plates break away from the mineralized fringes at variable distances from the gums. We proposed a model for estimating the effective pore size to determine whether flow scales with body length or prey size across species. We found that pore size is likely not a proxy for prey size but instead, may reflect changes in resistance through the filter that affect fluid flow., (© 2023 Wiley Periodicals LLC.)

Predation, combat, and the slings and arrows of an abrasive and high impact environment, represent just some of the biotic and abiotic stressors that fishes are armored against. The Pacific Spiny Lumpsucker (Eumicrotremus orbis) found in the subtidal of the Northern Pacific Ocean is a rotund fish covered with epidermal, cone-shaped, enamel odontodes. The Lumpsucker is a poor swimmer in the wave swept rocky intertidal, and this armor may be a lightweight solution to the problem of collisions with abiotic obstacles. We use micro-CT and scanning electron microscopy to reveal the morphology and ontogeny of the armor, and to quantify the amount of mineralization relative to the endoskeleton. The non-overlapping odontodes are organized into eight rows-six rows on the body, one row surrounding the eye, and one row underneath the chin. Odontodes start as a single, hooked cone; and they grow by the addition of cusps that accrete into a spiral. The mineral investment in armor compared to skeleton increases over ontogeny. Damage to the armor occurs both through passive abrasion and breakage from impact; and there is no evidence of replacement, or repair of damaged odontodes., (© 2022 Wiley Periodicals LLC.)

Ctenophores are descendants of an early branching basal metazoan lineage, which may have evolved neurons and muscles independently from other animals. Mnemiopsis is one of the important reference ctenophore species. However, little is known about its neuromuscular organization. Here, we mapped and tracked the development of the neural and muscular elements in the early hatching cydippid larvae, as well as adult Mnemiopsis leidyi. The overall development of the neuromuscular system in Mnemiopsis was very similar to Pleurobrachia bachei, although in Mnemiopsis the entire process occurred significantly faster. The subepithelial neural cells were observed immediately after hatching. This population consisted of a dozen of separated individual neurons with short neurites. In about 2 days, when their neurites grew significantly longer and connected to their neighbors, they began to form a canonical polygonal subepithelial network. Mesogleal neural elements prominent in all studied adult ctenophores were not detectable in Mnemiopsis larvae but were clearly labeled in closely related Lobata species Bolinopsis infundibulum. Hatched larvae also had putative mechanoreceptors with long stereocilia and approximately two dozen muscle cells. In adult Mnemiopsis, the feeding lobes and auricles contained two distinct populations of neurons and neural ensembles that were not observed in other ctenophore lineages and likely represented elaborate neuronal innovations characteristic for the clade Lobata and their lifestyles., (© 2021 Wiley Periodicals LLC.)

Many vertebrates are armored over all or part of their body. The armor may serve several functional roles including defense, offense, visual display, and signal of experience/capability. Different roles imply different tradeoffs; for example, defensive armor usually trades resistance to attack for maneuverability. The poachers (Agonidae), 47 species of scorpaeniform fishes, are a useful system for understanding the evolution and function of armor due to their variety and extent of armoring. Using publically available CT-scan data from 27 species in 16 of 21 genera of poachers we compared the armor to axial skeletal in the mid body region. The ratio of average armor density to average skeleton density ranged from 0.77 to 1.17. From a defensive point of view, the total investment in mineralization (volume * average density) is more interesting. There was 10 times the material invested in the armor as in the endoskeleton in some small, smooth plated species, like Aspidophoroides olrikii. At the low end, some visually arresting species like Percis japonica, had ratios as low as 2:1. We categorized the extent and type (impact vs. abrasion) in 34 Agonopsis vulsa across all 35+ plates in the eight rows along the body. The ventral rows show abrasive damage along the entire length of the fish that gets worse with age. Impact damage to head and tail plates gets more severe and occurs at higher rates with age. The observed damage rates and the large investment in mineralization of the armor suggest that it is not just for show, but is a functional defensive structure. We cannot say what the armor is defense against, but the abrasive damage on the ventrum implies their benthic lifestyle involves rubbing on the substrate. The impact damage could result from predatory attacks or from intraspecific combat., (© 2020 Wiley Periodicals LLC.)

Durophagous predators consume hard-shelled prey such as bivalves, gastropods, and large crustaceans, typically by crushing the mineralized exoskeleton. This is costly from the point of view of the bite forces involved, handling times, and the stresses inflicted on the predator's skeleton. It is not uncommon for durophagous taxa to display an ontogenetic shift from softer to harder prey items, implying that it is relatively difficult for smaller animals to consume shelled prey. Batoid fishes (rays, skates, sawfishes, and guitarfishes) have independently evolved durophagy multiple times, despite the challenges associated with crushing prey harder than their own cartilaginous skeleton. Potamotrygon leopoldi is a durophagous freshwater ray endemic to the Xingu River in Brazil, with a jaw morphology superficially similar to its distant durophagous marine relatives, eagle rays (e.g., Aetomylaeus, Aetobatus). We used second moment of area as a proxy for the ability to resist bending and analyzed the arrangement of the mineralized skeleton of the jaw of P. leopoldi over ontogeny using data from computed tomography (CT) scans. The jaws of P. leopoldi do not resist bending nearly as well as other durophagous elasmobranchs, and the jaws are stiffest nearest the joints rather than beneath the dentition. While second moment has similar material distribution over ontogeny, mineralization of the jaws under the teeth increases with age. Neonate rays have low jaw stiffness and poor mineralization, suggesting that P. leopoldi may not feed on hard-shelled prey early in life. These differences in the shape, stiffness and mineralization of the jaws of P. leopoldi compared to its durophagous relatives show there are several solutions to the problem of crushing shelled prey with a compliant skeleton., (© 2019 Wiley Periodicals, Inc.)

The Sarcastic Fringehead (Neoclinus blanchardi, Teleostei) exhibits an extreme version of a common aggressive display, the "gaping display," in which an open mouth is presented toward an opponent. Males of this species have extremely long jaws that extend posteriorly well past the posterior margin of the head and are flared laterally during the gaping display. In this study, we explored morphological traits related to this extraordinary display in this and related species of blennies. Morphological modifications include enlargement of the buccopalatal membrane, elongation of the maxilla via an uncalcified posterior extension, and evolution of a novel hinge between the anterior maxilla and lacrimal bones permitting lateral movement of the upper jaw. Geometric morphometry using the truss network system, thin-plate spline, and PCA of three closely related species of Neoclinus indicate that the elongate maxilla of N. blanchardi most likely evolved via acceleration (faster growth compared to outgroups) and hypermorphosis (continued growth to a larger body size), both forms of peramorphic heterochrony. Coloration and fluorescence of the buccopalatal membrane may also serve to amplify the extraordinary gaping display of the Sarcastic Fringehead., (© 2018 Wiley Periodicals, Inc.)

Members of the genus Paramoeba (including Neoparamoeba) (Amoebozoa) are single-celled eukaryotes of economic and ecological importance because of their association with disease in a variety of marine animals including fish, sea urchins, and lobster. Interestingly, they harbor a eukaryotic endosymbiont of kinetoplastid ancestry, Perkinsela sp. To investigate the complex relationship between Paramoeba spp. and Perkinsela sp., as well as the relationships between different Paramoeba species, molecular data was obtained for four novel isolates. We also acquired new data from the urchin pathogen P. invadens. Comprehensive molecular phylogenetic analyses were carried out using 33 newly obtained 18S rDNA sequences from the host amoebae and 16 new 18S rDNA sequences from their corresponding Perkinsela sp., together with all publicly available 18S molecular data. Intra-isolate 18S rDNA nucleotide diversity was found to be surprisingly high within the various species of Paramoeba, but relatively low within their Perkinsela sp. endosymbionts. 18S rDNA phylogenies and ParaFit co-evolution analysis revealed a high degree of congruence between the Paramoeba and Perkinsela sp. tree topologies, strongly suggesting that a single endosymbiotic event occurred in the common ancestor of known Paramoeba species, and that the endosymbionts have been inherited vertically ever since., (© 2017 The Author(s) Journal of Eukaryotic Microbiology © 2017 International Society of Protistologists.)

Background: Head or anterior body part regeneration is commonly associated with protostome, but not deuterostome invertebrates. However, it has been shown that the solitary hemichordate Ptychodera flava possesses the remarkable capacity to regenerate their entire nervous system, including their dorsal neural tube and their anterior head-like structure, or proboscis. Hemichordates, also known as acorn worms, are marine invertebrate deuterostomes that have retained chordate traits that were likely present in the deuterostome ancestor, placing these animals in a vital position to study regeneration and chordate evolution. All acorn worms have a tripartite body plan, with an anterior proboscis, middle collar region, and a posterior trunk. The collar houses a hollow, dorsal neural tube in ptychoderid hemichordates and numerous chordate genes involved in brain and spinal cord development are expressed in a similar anterior-posterior spatial arrangement along the body axis., Results: We have examined anterior regeneration in the hemichordate Ptychodera flava and report the spatial and temporal morphological changes that occur. Additionally, we have sequenced, assembled, and analyzed the transcriptome for eight stages of regenerating P. flava, revealing significant differential gene expression between regenerating and control animals., Conclusions: Importantly, we have uncovered developmental steps that are regeneration-specific and do not strictly follow the embryonic program. Developmental Dynamics 245:1159-1175, 2016. © 2016 The Authors. Developmental Dynamics published by Wiley Periodicals, Inc. on behalf of American Association of Anatomists., (© 2016 The Authors. Developmental Dynamics published by Wiley Periodicals, Inc. on behalf of American Association of Anatomists.)

Premise of the Study: Morphology and material properties are the main components of the mechanical design of organisms, with species groups developing different optimization strategies in the context of their physical environment. For intertidal and subtidal seaweeds, possessing highly flexible and extensible tissues allows individuals to bend and reconfigure in flow, thereby reducing drag. Previous research has shown that aging may compromise these qualities. Tissue age increases with distance from the blade's meristem, which differs in its position on kelps and red algae. Here, we assess whether longitudinal patterns of blade material properties differ between these two algal groups according to tissue age., Methods: We performed tensile tests on tissues samples excised from various positions along the extent of blades in nine kelp species (basal growth) and 15 species of red algae (apical growth)., Key Results: We found that older tissues were less flexible and extensible than younger tissues in all species tested. As predicted, tissue near the basal meristem in kelp was more flexible and extensible than older tissue at the blade's distal end. The opposite pattern was observed for red algae, with the most flexible and extensible tissues found near the apical meristem at the distal ends of blades., Conclusions: We propose that divergent patterns in the distribution of material properties along blades may have different consequences for the performance of kelps and red algae. The positioning of younger tissues at the blade base for kelps may enable these species to attain larger body sizes in wave-swept habitats., (© 2015 Botanical Society of America.)

Three lineages of cartilaginous fishes have independently evolved filter feeding (Lamniformes: Megachasma and Cetorhinus, Orectolobiformes: Rhincodon, and Mobulidae: Manta and Mobula); and the structure of the branchial filters is different in each group. The filter in Rhincodon typus has been described; species within the Lamniformes have simple filamentous filters, but the anatomy and ultrastructure of the branchial filter in the mobulid rays varies and is of functional interest. In most fishes, branchial gill rakers are elongated structures located along the anterior ceratobranchial and/or epibranchial arches; however, mobulid gill rakers are highly modified, flattened, lobe-like structures located on the anterior and posterior epibranchial elements as well as the ceratobranchials. The ultrastructure of the filter lobes can be smooth or covered by a layer of microcilia, and some are denticulated along the dorsal and ventral lobe surface. Flow through the mobulid oropharyngeal cavity differs from other filter-feeding fishes in that water must rapidly deviate from the free stream direction. There is an abrupt 90° turn from the initial inflowing path to move through the laterally directed branchial filter pores, over the gill tissue, and out the ventrally located gill slits. The deviation in the flow must result in tangential shearing stress across the filter surface. This implies that mobulids can use cross-flow filtration in which this shearing force serves as a mechanism to resuspend food particles initially caught by sieving or another capture mode. These particles will be transported by the cross filter flow toward the esophagus. We propose that species with cilia on the rakers augment the shear mediated movement of particles along the filter with ciliary transport., (Copyright © 2013 Wiley Periodicals, Inc.)

The process of spermiation in the ratfish Hydrolagus colliei is described and compared with that in mammals and amphibians. Spermiation in this species involves prior fluid space expansion both within the apical parts of the Sertoli cytoplasm and in the spaces between Sertoli cells and spermatids. The apical ends of Sertoli cells fragment, including the parts immediately around the spermatid acrosomes. Intercellular material between the spermatid tips and the Sertoli cells dissolves. Concurrently an opening from the seminiferous follicle into the efferent ductule is made by means of changes in cell shape and separation of Sertoli cells and efferent ductule cells that adhere to each other up to the time of sperm release., (Copyright © 1990 Wiley-Liss, Inc.)

The ultrastructure of the stylets produced by nine species of nemerteans has been examined by scanning electron microscopy (S.E.M.) and polarized light microscopy. Stylets are solid, nail-shaped structures that typically reach lengths of 50-200 μm. Each stylet is composed of a centrally located organic matrix surrounded by an inorganic cortex that contains calcium and phosphorus. When viewed at high magnifications, fine granules can be seen throughout the organic matrix, and the cortex appears to be composed of densely packed homo-geneous material. Fractured specimens and whole matrices isolated from decalcified stylets reveal a close correspondence between the shape of the organic matrix and that of the surrounding cortex. This similarity in morphology suggests that the organic matrix serves as a template during calcification of the stylet. The fact that abundant material can be seen in the core of incinerated stylets, and in the central region of stylets that had been soaked for several hours in sodium hypochlorite, supports the hypothesis that the organic matrix is also highly calcified. Polarization microscopy of nemertean stylets indicates that they are composed of a crystalline, rather than amorphous, form of calcium phosphate. The probable organization of the calcium phosphate crystals is discussed., (Copyright © 1983 Wiley-Liss, Inc.)

The morphology of the newly hatched larva of Carcinonemertes epialti Coe has been examined by light and electron microscopy. The newly hatched larva is covered with cilia and measures about 110 μm in length. Four types of epidermal cells are recognizable: (1) Multiciliated cells, (2) vacuolated cells, (3) mucous cells, and (4) "knob cells". The knob cells protrude from the posterior end of the larva and contain granules and bundles of microfilaments. The gut is incomplete and is located ventral to the bipartite proboscis. A bilobed brain and two subepidermal ocelli are found in the anterior end of the larva. The anterior and posterior cirri are composed of long, tightly appressed cilia that arise from an invagination of the epidermis at each end of the larva. The anterior cirrus is surrounded by two types of glandular cells. It is proposed that the knob cells have a role in larval attachment, combining the functions of the adhesive cells and anchor cells described in the duo-gland system of turbellarians. The cirri are believed to be larval sensory structures that function in substrate selection. Histological and ultrastructural observations suggest that the larvae of Carcinonemertes are relatively long lived and develop into juveniles without a drastic metamorphosis., (Copyright © 1981 Wiley-Liss, Inc.)

The ultrastructure of the "spiny" surface of Tealia crassicornis eggs is examined in detail by scanning and transmission electron microscopy in order to understand its function. Long microvilli are clustered together in spiral aggregates of 50-75 microvilli called "spires." There are about 15,000 spires per egg. Dense bundles of microfilaments making up the cores of these microvilli are shown to be composed of actin by staining with the fluorescent dye nitrobenzoxadiazole (NBD)-phallacidin. It is postulated that the bundles of actin and the spires of microvilli are stiff and provide reinforcement to the egg surface. Such postulated properties would provide physical protection for these large eggs which, unlike the eggs of most invertebrates, appear to lack all extracellular investing coats., (Copyright © 1982 Wiley-Liss, Inc.)

The nemertean Paranemertes peregrina uses its long, eversible proboscis to capture nereid polychaetes. During prey capture, the glandular epithelium of the everted proboscis secretes a sticky venom that contains a potent neurotoxin. In this study, the ultrastructure of the venom-producing epithelium is examined before and after capture of nereid prey. Four types of cells can be distinguished in the glandular epithelium, based on the shape and staining properties of their secretory products. The cells contain fusiform or coalescent types of rhabdoids, acidophilic granules, or flocculent material. All four cell types occur in the anterior part of the proboscis, but only cells with acidophilic granules have been observed in the posterior chamber. The glandular epithelium of the anterior chamber secretes sheets of venomous mucus that are composed of the four types of secretory products fused together, while the epithelial cells of the posterior chamber produce a granulated discharge. Cells that contain flocculent material also occur in the epidermis of P. peregrina. The flocculent material is believed to contain the toxic component of the venom, as its distribution in the anterior proboscis chamber and body wall correlates with previous reports of toxin concentrations that were derived from assays of tissue extracts., (Copyright © 1983 Wiley-Liss, Inc.)

The ultrastructure of cellular organelles involved in stylet formation is examined in six species of nemertean worms by transmission electron microscopy (TEM). Stylets are nail-shaped structures containing calcium phosphate that are assembled intracellularly in large uninucleate cells, called styletocytes. Each stylet develops within a membrane-bound vacuole in the styletocyte cytoplasm. Well developed arrays of Golgi bodies are typically found in the vicinity of developing stylet vacuoles, and fully formed vacuoles are filled with PAS+ material that appears to be derived from the smooth endoplasmic reticulum. At the onset of styletogenesis, a conical sliver of organic material differentiates on the inner surface of the vacuolar membrane. This material displays a species-specific banding pattern in decalcified sections, and apparently acts as a template during calcification of the stylet shaft. After the organic core of the shaft is formed, mitochondria aggregate around the stylet vacuole and presumably help accumulate the calcium used in mineralization of the stylet. A knob-shaped proximal piece is subsequently assembled on the base of the shaft. The proximal piece contains a nonbanded matrix and has electron-dense material at its surface that may help in correctly orienting this region toward the basis during replacement of the central stylet., (Copyright © 1984 Wiley-Liss, Inc.)