Versatile ciliary behaviour in capture of particles by the bryozoan cyphonautes larva

Cyphonautes larvae of a bryozoan, Membranipora membranacea, used several ciliary mechanisms to capture algal cells upstream from the lateral band of cilia that produces a feeding current. (1) Lateral cilia changed beat and a backcurrent occurred at the time and place that particles were retained. (2) Algal cells were sieved and held stationary at the upstream (frontal) side of a row of laterofrontal cilia that were not beating. (3) Localized extension of cilia toward the inhalant chamber, coincident with particle captures, indicated that laterofrontal cilia flick toward the inhalant chamber. These flicks may aid transport of captured particles toward the mouth. Thus my earlier report that larvae only sieve, in contrast to the adults (which have an active ciliary response) was in error. The similar ciliary bands in adult and larval bryozoans and in other lophophorates (brachiopods, and phoronids) suggest that these animals share a core repertoire of ciliary behaviours in the capture and concentration of suspended food particles.     

Ciliary filter-feeding structures in adult and larval gymnolaemate bryozoans

Abstract. Ciliary filter-feeding structures of gymnolaemate bryozoans—adults of Flustrellidra hispida and Alcyonidium gelatinosum , larvae of Membranipora sp.—were studied with SEM. In F. hispida and A. gelatinosum , the distal part of each tentacle has a straight row of stiff laterofrontal cilia which carry out "ciliary sieving" to capture suspended food particles that are subsequently transported downward towards the mouth by tentacle flicking; both structure and function resemble those of stenolaemate tentacles. The proximal part of the tentacle and of the ciliary ridge of a cyphonautes larva have strikingly similar structures, except that the laterofrontal cells are monociliate in the adults and biciliate in the larvae. The laterofrontal cells of the tentacles are arranged in a zigzag row and their cilia form two parallel rows, a frontal and a lateral row. The latter probably forms the sieve of stiff filter cilia in front of the water-pumping lateral cilia, whereas the frontal row appears to be held close to the frontal ciliary band of the tentacle. The biciliate laterofrontal cells of the cyphonautes larva have the cilia arranged in similar rows. The detailed morphological similarities between the ciliary bands of adult and larval filtering structures suggest that the feeding mechanisms are similar, contrary to what has been previously thought.     

Ciliary feeding structures and particle capture mechanism in the freshwater bryozoan Plumatella repens (Phylactolaemata)

Abstract. In contrast to marine bryozoans, the lophophore structure and the ciliary filter‐feeding mechanism in freshwater bryozoans have so far been only poorly described. Specimens of the phylactolaemate bryozoan Plumatella repens were studied to clarify the tentacular ciliary structures and the particle capture mechanism. Scanning electron microscopy revealed that the tentacles of the lophophore have a frontal band of densely packed cilia, and on each side a zigzag row of laterofrontal cilia and a band of lateral cilia. Phalloidin‐linked fluorescent dye showed no sign of muscular tissue within the tentacles. Video microscopy was used to describe basic characteristics of particle capture. Suspended particles in the incoming water flow, set up by the lateral ‘pump’ cilia on the tentacles, approach the tentacles with a velocity of 1–2 mm s^‐1. Near the tentacles, the particles are stopped by the stiff sensory laterofrontal cilia acting as a mechanical sieve, as previously seen in marine bryozoans. The particle capture mechanism suggested is based on the assumed ability of the sensory stiff laterofrontal cilia to be triggered by the deflection caused by the drag force of the through‐flowing water on a captured food particle. Thus, when a particle is stopped by the laterofrontal cilia, the otherwise stiff cilia are presumably triggered to make an inward flick which brings the restrained particle back into the downward directed main current, possibly to be captured again further down in the lophophore before being carried to the mouth via the food groove. No tentacle flicks and no transport of captured particles on the frontal side of the tentacles were observed. The velocity of the metachronal wave of the water‐pumping lateral cilia was measured to be ～0.2 mm s^‐1, the wavelength was ～7 μm, and hence the ciliary beat frequency estimated to be ～30 Hz (～20 °C). The filter feeding process in P. repens reported here resembles the ciliary sieving process described for marine bryozoans in recent years, although no tentacle flicks were observed in P. repens. The phylogenetic position of the phylactolaemates is discussed in the light of these findings.     

Cinefilms of particle capture by an induced local change of beat of lateral cilia of a bryozoan

A local disruption of the metachronal wave always accompanies capture of algal cells by tentacles of Flustrellidra hispida (Fabricius). Beat changes for ≈0.2 s over ≈100μm of the ciliated band during capture of a 10-μm particle. The halted parcel of water is therefore larger than the particle of food but much smaller than the flow that continues past the tentacles elsewhere. These events are consistent with the hypothesis that an induced local reversal of beat concentrates particles for those suspension feeders that retain particles upstream from a band of simple cilia (adults or larvae of bryozoans, brachiopods, phoronids, hemichordates, and echinoderms). These events are not explained by other hypotheses that have been advanced for concentration of particles by these suspension feeders. Aerosol filtration models of direct interception are not applicable to this type of ciliary suspension feeder because retention depends on the magnitude of a stimulus and response to it. The stimulus will not be the same function of diameter of the food particle, and response is unlikely below a threshold stimulus.     

Studies on the gill ciliation of the American oyster Crassostrea virginica (Gmelin)

Latero-frontal, para-latero-frontal, and frontal ciliary tracts on the gill filaments of Crassostrea virginica (Gmelin) were studied with light microscopy and scanning electron microscopy. Latero-frontal cirri are complex structures composed of varying numbers of paired cilia. The multiple pairs of cilia which constitute a single cirrus are closely appressed for a portion of their length; they then branch laterally from the central axis in a plume-like fashion. Latero-frontal cirri of adjacent gill filaments create a filtration sieve which should be capable of retaining particles smaller than 1 μm in diameter. Para-latero-frontal cilia are short, closely spaced cilia arranged as a staggered row along the frontal side of each tract of latero-frontal cirri. Latero-frontal cirri and para-latero-frontal cilia occur on ordinary, principal, and transitional gill filaments. Frontal ciliary tracts of ordinary filaments are divided into a central, ventrally directed coarse tract, flanked on either side by a dorsally directed fine ciliary tract. The coarse tract is covered by cirri which are comprised of five to eight cilia, while the fine frontal tracts are made up of individually functioning cilia. The frontal ciliary tracts of principal and transitional filaments bear only dorsally directed fine cilia. The unique direction of effective beat of the coarse frontal cirri of ordinary filaments, in combination with the action of fine frontal cilia and the strategic location of mucus producing cells, is used to describe a possible mechanism for the sorting of filtered particles.     

Mechanical sensitivity and cell coupling in the ciliated epithelial cells ofMytilus edulis gill

Summary Transmission electron microscopy has not provided strong evidence for gap junctions inMytilus edulis gill tissue, in spite of extensive physiological evidence for coupled ciliary arrest in lateral cells and coupled activation in abfrontal cells. To investigate the kinds and relative distribution of cell junctions and also to determine whether ciliary membrane particle differences exist in these two types of oppositely mechanically sensitive cells, we analyzed the structure of these and two other ciliated cell types (frontal and laterofrontal) by freeze-fracture replication. Gap junctions occur in all four ciliated cell types, but they are relatively small and of variable morphology, often consisting of elongate, winding complexes of membrane particles. Statistically, such structures rarely would be recognized as gap junctions in thin sections. Gap junctions appear to be most abundant between the highly coupled abfrontal cells, minimal between laterofrontal cells, and not evident in the epithelial cells that separate coupled ciliated cell types. The ciliary necklaces of the mechanically activated abfrontal cilia are typically 4- or 5-stranded while those of the remaining three cell types are mainly 3-stranded. In developing gill tips, ciliated cells have abundant gap junctions and newly formed cilia have a full complement of necklace particles. Nascent lateral cilia are not mechanically sensitive, indicating that the acquisition of mechanosensitivity does not correlate with the presence of ciliary necklace or other membrane particles. Lateral and laterofrontal cells become sensitive to neurotransmitters soon after the appearance of the latter during development, but mechanosensitivity of both lateral and abfrontal cells arises substantially later.     

The mussel filter–pump – present understanding,with a re‐examination of gill preparations

Filter feeding in mussels is a secondary adaptation where the gills have become W‐shaped and greatly enlarged, acting as the mussel filter–pump. Water pumping and particle capture in the blue mussel, Mytilus edulis, have been studied over many years. Here, we give a short status of the present understanding of ciliary structure and function of the mussel filter–pump, supplemented with new photo‐microscope and scanning electron microscopy (SEM) pictures of gill preparations. Pumping rate (filtration) and pressure to maintain flow have been extensively studied so the power delivered by the mussel pump to the water flow is known (1.1% of total respiratory power), but the actual cost based on gill respiration is much higher (19%), implying that the cost of maintaining of the large gill pump is considerable and that only relatively little energy can be saved by stopping or reducing the activity of the water‐pumping cilia so that continuous feeding with a ‘minimal scaled’ pump is cheaper than discontinuous feeding with a correspondingly larger pump. According to the present view, the pump proper is the beating lateral cilia (lc) on the gill filaments and particle capture is accomplished by the action of laterofrontal cirri (lfc) transferring particles from the main water current to the frontal gill filament currents driven by frontal cilia (fc). Unexplained aspects include retention efficiency according to particle size and the role of pro‐laterofrontal cilia (p‐lfc) placed between the lfc and fc. The structure of cilia and the mode of ciliary beating have been re‐examined in this study by new high‐resolution light and scanning electron microscopy of isolated gill preparations exposed to serotonin (5‐HT) stimulation which can activate the lc and lfc at low concentrations (10^?6 M), but removes the lfc from the interfilament canals at higher concentrations (10^?5 M).     

Time and extent of ciliary response to particles in a non-filtering feeding mechanism

Mechanisms of suspension feeding are usually described by the physics of inanimate filters. High-speed videorecordings in this study demonstrated that sea urchin larvae concentrate particles without filtration. They actively captured individual particles. At most times and places, the effective strokes of the swimming/feeding ciliary band were away from the circumoral field. Cilia of this band responded to particles by a reversal of beat that redirected the particle toward the circumoral field. A change of beat occurred along approximately 80 micro m of ciliary band during particle capture. Cilia responded 0.02 to 0.06 s after the particle was within reach of effective strokes and reversed beat, usually for about 0.1 to 0.2 s. The whole event (disruption of forward beat) generally lasted between 0.13 and 0.5 s. These observations imply reversed movement of a parcel of water much larger than the included captured particle, but particles are nevertheless greatly concentrated because water is directed toward the circumoral field only when and where a particle is sensed. Thus most of the concentration of particles occurs by a temporarily and locally redirected current, without filtration, and size and quality of particles captured depends on sensory capabilities, not the mechanics of filtration.     

Opposed ciliary bands in the feeding larvae of sabellariid annelids

The larvae of marine annelids capture food using an unusual diversity of suspension-feeding mechanisms. Many of the feeding mechanisms of larval annelids are poorly known despite the abundance and ecological significance of both larvae and adults of some annelid taxa. Here we show that larvae of two species of sabellariid annelids, Sabellaria cementarium and Phragmatopoma californica, bear prototrochal and metatrochal cilia that beat in opposition to each other. For larvae of S. cementarium, we provide evidence that these opposed bands of cilia are used to capture suspended particles. In video recordings, captured particles were overtaken by a prototrochal cilium and then moved with the cilium to the food groove, a band of cilia between the prototroch and metatroch. They were then transported by cilia of the food groove to the mouth. Lengths of the prototrochal cilia, lengths of the prototrochal ciliary band, size range of the particles captured, and estimated rates of clearance increased with larval age and body size. Confirmation of the presence of opposed bands in larvae of sabellariids extends their known occurrence in the annelids to members of 10 families. Opposed bands in these different taxa differ in the arrangements and spacing of prototrochal and metatrochal cilia, and in whether they are used in combination with other feeding mechanisms. Opposed bands appear to be particularly widespread among the larvae of sabellidan annelids (a clade that includes sabellariids, sabellids, and serpulids), even in some species whose larvae do not feed. A parsimony analysis suggests that opposed bands are ancestral in this clade of annelids.     

Laser-induced spreading arrest of Mytilus gill cilia

Using a "slit camera" recording technique, we have examined the effects of local laser irradiation of cilia of the gill epithelium of Mytilus edulis. The laser produces a lesion which interrupts epithelial integrity. In artificial sea water that contains high K+ or is effectively Ca++ free, metachronism of the lateral cilia continues to either side of the lesion with only minor perturbations in frequency synchronization and wave velocity, such as would be expected if metachronal wave coordination is mechanical. However, in normal sea water and other appropriate ionic conditions (i.e., where Ca++ concentration is elevated), in addition to local damage, the laser induces distinct arrest responses of the lateral cilia. Arrest is not mechanically coordinated, since cilia stop in sequence depending on stroke position as well as distance from the lesion. The velocity of arrest under standard conditions is about 3 mm/s, several orders of magnitude faster than spreading velocities associated with diffusion of materials from the injured region. Two responses can be distinguished on the basis of the kinetics of recovery of the arrested regions. These are (a) a nondecremental response that resembles spontaneous ciliary stoppage in the gills, and (b) a decremental response, where arrest nearer the stimulus point is much longer lasting. The slower recovery is often periodic, with a step size approximating lateral cell length. Arrest responses with altered kinetics also occur in laterofrontal cilia. The responses of Mytilus lateral cilia resemble the spreading ciliary arrest seen in Elliptio and arrest induced by electrical and other stimuli, and the decremental response may depend upon electrotonic spread of potential change produced at the stimulus site. If this were coupled to transient changes in Ca++ permeability of the cell membrane, a local rise in Ca++ concentration might inhibit ciliary beat at a sensitive point in the stroke cycle to produce the observed arrest.     

Larval Feeding in Echinoderms

In all four types of feeding echinoderm larvae, particles areretained upstream from the ciliated band, probably by an inducedlocal reversal of ciliary beat. Comparative studies and theoreticalconsiderations suggest that increasing the length of the bandmay be the only means of increasing the rate at which wateris processed for paniculate food. This would account for thelong looping band and late development of adult structures inechinoderm larvae. Estimates of the minimum food requirementsof early echinplutei and of uptake of amino acids by embryosare calculated. Various means of rejecting particles are describedand observations related to nervous control of feeding and rejectionare discussed. The possible disadvantages of larvae developingfrom smaller but more numerous eggs are discussed. It is arguedthat loss of a feeding larval stage is usually an irreversibleevolutionary change in echinoderms, and some general implicationsof the irreversibility of such a change in life history arementioned.     

Visualization of calcium transients controlling orientation of ciliary beat

To image changes in intraciliary Ca controlling ciliary motility, we microinjected Ca Green dextran, a visible wavelength fluorescent Ca indicator, into eggs or two cell stages of the ctenophore Mnemiopsis leidyi. The embryos developed normally into free-swimming, approximately 0.5 mm cydippid larvae with cells and ciliary comb plates (approximately 100 microns long) loaded with the dye. Comb plates of larvae, like those of adult ctenophores, undergo spontaneous or electrically stimulated reversal of beat direction, triggered by Ca influx through voltage-sensitive Ca channels. Comb plates of larvae loaded with Ca Green dextran emit spontaneous or electrically stimulated fluorescent flashes along the entire length of their cilia, correlated with ciliary reversal. Fluorescence intensity peaks rapidly (34-50 ms), then slowly falls to resting level in approximately 1 s. Electrically stimulated Ca Green emissions often increase in steps to a maximum value near the end of the stimulus pulse train, and slowly decline in 1-2 s. In both spontaneous and electrically stimulated flashes, measurements at multiple sites along a single comb plate show that Ca Green fluorescence rises within 17 ms (1 video field) and to a similar relative extent above resting level from base to tip of the cilia. The decline of fluorescence intensity also begins simultaneously and proceeds at similar rates along the ciliary length. Ca-free sea water reversibly abolishes spontaneous and electrically stimulated Ca Green ciliary emissions as well as reversed beating. Calculations of Ca diffusion from the ciliary base show that Ca must enter the comb plate along the entire length of the ciliary membranes. The voltage-dependent Ca channels mediating changes in beat direction are therefore distributed over the length of the comb plate cilia. The observed rapid and virtually instantaneous Ca signal throughout the intraciliary space may be necessary for reprogramming the pattern of dynein activity responsible for reorientation of the ciliary beat cycle.     

Swimming Speed of Larval Snail Does Not Correlate with Size and Ciliary Beat Frequency

Many marine invertebrates have planktonic larvae with cilia used for both propulsion and capturing of food particles. Hence, changes in ciliary activity have implications for larval nutrition and ability to navigate the water column, which in turn affect survival and dispersal. Using high-speed high-resolution microvideography, we examined the relationship between swimming speed, velar arrangements, and ciliary beat frequency of freely swimming veliger larvae of the gastropod Crepidula fornicata over the course of larval development. Average swimming speed was greatest 6 days post hatching, suggesting a reduction in swimming speed towards settlement. At a given age, veliger larvae have highly variable speeds (0.8–4 body lengths s⁻¹) that are independent of shell size. Contrary to the hypothesis that an increase in ciliary beat frequency increases work done, and therefore speed, there was no significant correlation between swimming speed and ciliary beat frequency. Instead, there are significant correlations between swimming speed and visible area of the velar lobe, and distance between centroids of velum and larval shell. These observations suggest an alternative hypothesis that, instead of modifying ciliary beat frequency, larval C. fornicata modify swimming through adjustment of velum extension or orientation. The ability to adjust velum position could influence particle capture efficiency and fluid disturbance and help promote survival in the plankton.     

Large‐scale ciliary reversal mediates capture of individual algal prey by Müller's larva

We documented capture of microalgal prey by several species of wild‐caught Müller's larvae of polyclad flatworms. To our knowledge, this is the first direct observation of feeding mechanism in this classical larval type. High‐speed video recordings showed that virtually all captures were mediated by large‐scale transient ciliary reversal over one or more portions of the main ciliary band corresponding to individual lobes or tentacles. Local ciliary beat reversals altered near‐field flow to suck parcels of food‐containing water mouthward. Many capture episodes entailed sufficient coordinated flow disruption that these compact‐bodied larvae tumbled dramatically. Similar behaviors were recorded in at least four distinct species, one of which corresponds to the ascidian‐eating polyclad Pseudoceros canadensis.     

Numerical simulation of flow fields and particle trajectories in ciliary suspension feeding

A model describing the ciliary driven flow and motion of suspended particles in downstream suspension feeders is developed. The quasi-steady Stokes equations for creeping flow are solved numerically in an unbounded fluid domain around cylindrical bodies using a boundary integral formulation. The time-dependent flow is approximated with a continuous sequence of steady state creeping flow fields, where metachronously beating ciliary bands are modelled by linear combinations of singularity solutions to the Stokes equations. Generally, the computed flow fields can be divided into an unsteady region close to the driving ciliary bands and a steady region covering the remaining fluid domain. The size of the unsteady region appears to be comparable to the metachronal wavelength of the ciliary band. A systematic investigation is performed of trajectories of infinitely small (fluid) particles in the simulated unsteady ciliary driven flow. A fraction of particles appear to follow trajectories, that resemble experimentally observed particle capture events in the downstream feeding system of the polycheate Sabella penicillus, indicating that particles can be captured by ciliary systems without mechanical contact between particle and cilia. A local capture efficiency is defined and its value computed for various values of beat frequencies and other parameters. The results indicate that the simulated particle capture process is most effective when the flow field oscillates within timescales comparable to transit timescales of suspended particles passing the unsteady region near the ciliary bands. However, the computed retention efficiencies are found to be much lower than those obtained experimentally.     

Ciliary reversal without rotation of axonemal structures in ctenophore comb plates

We have used a newly discovered reversal response of ctenophore comb plates to investigate the structural mechanisms controlling the direction of ciliary bending. High K+ concentrations cause cydippid larvae of the ctenophore Pleurobrachia to swim backward. High-speed cine films of backward-swimming animals show a 180 degree reversal in beat direction of the comb plates. Ion substitution and blocking experiments with artificial seawaters demonstrate that ciliary reversal is a Ca++-dependent response. Comb plate cilia possess unique morphological markers for numbering specific outer-doublet microtubules and identifying the sidedness of the central pair. Comb plates of forward- and backward-swimming ctenophores were frozen in different stages of the beat cycle by an "instantaneous fixation" method. Analysis of transverse and longitudinal sections of instantaneously fixed cilia showed that the assembly of outer doublets does not twist during ciliary reversal. This directly confirms the existence of radial switching mechanism regulating the sequence of active sliding on opposite sides of the axoneme. We also found that the axis of the central pair always remains perpendicular to the plane of bending; more importantly, the ultrastructural marker showed that the central pair does not rotate during a 180 degree reversal in beat direction. Thus, the orientation of the central pair does not control the direction of ciliary bending (i.e., the pattern of active sliding around the axoneme). We discuss the validity of this finding for three-dimensional as well as two-dimensional ciliary beat cycles and conclude that models of central-pair function based on correlative data alone must now be re-examined in light of these new findings on causal relations.     

CILIARY MEMBRANE DIFFERENTIATIONS IN TETRAHYMENA PYRIFORMIS : Tetrahymena Has Four Types of Cilia

We have examined thin sections and replicas of freeze-fractured cilia of Tetrahymena pyriformis. The ciliary necklace located at the base of all freeze-fractured oral and somatic cilia has been studied in thin sections. Since electron-dense linkers have been found to connect both microtubule doublets and triplets to the ciliary membrane at the level of the necklace, the linkers and the associated necklace seem to be related to the transition region between the doublets and triplets of a cilium. Plaque structures, consisting of small rectangular patches of particles located distal to the ciliary necklace, are found in strain GL, but are absent in other strains examined in this study. In freeze-cleaved material, additional structural differentiations are observed in the distal region of the ciliary membranes of somatic and oral cilia. Somatic cilia contain many randomly distributed particles within their membrane. Oral cilia can be divided into three categories on the basis of the morphology of their freeze-fractured membranes: (a) undifferentiated cilia with very few randomly distributed particles: (b) cilia with particles arranged in parallel longitudinal rows spaced at intervals of 810–1080 Å that are located on one side of the cilium; and (c) cilia with patches of particles arranged in short rows oriented obliquely to the main axis of the cilium. The latter particles, found on one side of the cilium, seem to serve as attachment sites for bristles 375–750 Å long and 100 Å wide which extend into the surrounding medium. The particles with bristles are located at the tips of cilia in the outermost membranelle and may be used to detect food particles and/or to modify currents in the oral region so that food particles are propelled more efficiently into the buccal cavity. Examination of thin-sectioned material indicates that the particles in oral cilia which form the longitudinal rows could be linked to microtubule doublets. Linkage between microtubule doublets and adjacent membrane areas on one side of the cilium could modify the form of ciliary beat by restricting the sliding of the microtubules. It is suggested that membrane-microtubule interactions may form the basis for the various forms of ciliary beat observed in different organisms.     

Determination of ciliary polarity precedes differentiation in the epithelial cells of quail oviduct.

In quail oviduct epithelium, as in all metazoan and protozoan ciliated cells, cilia beat in a coordinated cycle. They are arranged in a polarized pattern oriented according to the anteroposterior axis of the oviduct and are most likely responsible for transport of the ovum and egg white proteins from the infundibulum toward the uterus. Orientation of ciliary beating is related to that of the basal bodies, indicated by the location of the lateral basal foot, which points in the direction of the active stroke of ciliary beating. This arrangement of the ciliary cortex occurs as the ultimate step in ciliogenesis and following the oviduct development. Cilia first develop in a random orientation and reorient later, simultaneously with the development of the cortical cytoskeleton. In order to know when the final orientation of basal bodies and cilia is determined in the course of oviduct development, microsurgical reversal of a segment of the immature oviduct was performed. Then, after hormone-induced development and ciliogenesis, ciliary orientation was examined in the inverted segment and in normal parts of the ciliated epithelium. In the inverted segment, orientation was reversed, as shown by a video recording of the direction of effective flow produced by beating cilia, by the three-dimensional bending forms of cilia immobilized during the beating cycle and screened by scanning electron microscopy, and by the position of basal body appendages as seen in thin sections by transmission electron microscopy. These results demonstrate that basal body and ciliary orientation are irreversibly determined prior to development by an endogenous signal present early in the cells of the immature oviduct, transmitted to daughter cells during the proliferative phase and expressed at the end of ciliogenesis.     

Cilia and the life of ctenophores

Ctenophores, or comb jellies, are a distinct phylum of marine zooplankton with eight meridional rows of giant locomotory comb plates. Comb plates are the largest ciliary structures known, and provide unique experimental advantages for investigating the biology of cilia. Here, I review published and unpublished work on how ctenophores exploit both motile and sensory functions of cilia for much of their behavior. The long‐standing problem of ciliary coordination has been elucidated by experiments on a variety of ctenophores. The statocyst of ctenophores is an example of how mechanosensory properties of motile cilia orient animals to the direction of gravity. Excitation or inhibition of comb row beating provides adaptive locomotory responses, and global reversal of beat direction causes escape swimming. The diverse types of prey and feeding mechanisms of ctenophores are related to radiation in body form and morphology. The cydippid Pleurobrachia catches copepods on tentacles and undergoes unilateral ciliary reversal to sweep prey into its mouth. Mnemiopsis uses broad muscular lobes and ciliated auricles to capture and ingest prey. Beroë has giant smooth muscles and toothed macrocilia to rapidly engulf or bite through ctenophore prey, and uses reversible tissue adhesion to keep its mouth closed while swimming. Ciliary motor responses are calcium‐dependent, triggered by voltage‐activated calcium channels located along the length (reversed beating) or at the base (activation of beating) of ciliary membranes. Ciliary and muscular responses to stimuli are regulated by epithelial and mesogleal nerve nets with ultrastructurally identifiable synapses onto effector cells. Post‐embryonic patterns of comb row development in larval and adult stages are described and compared with regeneration of comb plates after surgical removal. Truly, cilia and ctenophores, like love and marriage, go together like a horse and carriage.     

Opposed bands of cilia in the nonfeeding larvae of the serpulid annelid Salmacina tribranchiata

The nonfeeding planktonic larvae of marine invertebrates typically lack larval feeding structures. One puzzling exception to this generalization is the annelid clade Sabellidae, in which nonfeeding larvae possess ciliary bands (specifically, food groove and metatroch) that, to the best of our knowledge, have no function other than in feeding. Nishi and Yamasu (1992b, Bulletin of the College of Sciences, University of the Ryukyus, 54 , 107–121) published a scanning electron micrograph showing that nonfeeding larvae of the serpulid annelid Salmacina dysteri also possess food groove and metatrochal cilia. Here I demonstrate that nonfeeding larvae of Salmacina tribranchiata also bear ciliary bands identifiable as food groove and metatroch by position. High‐speed video of ciliary beat patterns shows that, together with the prototrochal cilia, these bands function in an opposed band system. The presence of feeding structures in nonfeeding annelid larvae is thus more widely distributed than previously recognized. The presence of feeding structures may make evolutionary transitions to planktotrophy more likely, and may underlie an inferred origin of larval feeding in the common ancestor of one of the two major clades of serpulid annelids, Serpulinae.