Behavioral responses of a larval crustacean to hydrostatic pressure: Rhithropanopeus harrisii (Brachyura: Xanthidae)

Responses of the four zoeal stages of the crab Rhithropanopeus harrisii (Gould) to step and continuous changes in hydrostatic pressure were analyzed with a video system. Crabs were collected from the Neuse River estuary (North Carolina, USA) from June to August, 1987. The lower thresholds for step increases and decreases in pressure were 3 and 8 to 10 mbar, respectively. There was little change in sensitivity with zoeal development. Tests of larval responses in a light field that simulated the underwater angular light distribution indicated positive phototaxis does not occur upon pressure changes. In darkness, rates of pressure increase at and above 0.175 mbar s^-1 induced high barokinesis and negative geotaxis in all but Stage IV zoeae, which had a threshold of 1.19 mbar s^-1. Since larval sinking and descent swimming speeds exceed these threshold rates, larvae can move rapidly enough to produce suprathreshold changes in pressure which evoke behavioral responses. Slow rates of pressure decrease induced passive sinking while rapid rates caused an active ascent. This ascent response upon a pressure decrease is unreported among crustaceans, and is hypothesized to function for avoidance of feeding and respiratory currents of benthic invertebrates. The descent response occurs in all zoeal stages, except IV, at rates of pressure decrease (0.4 to 0.71 mbar s^-1) that are within the range of ascent swimming speeds. These results support Sulkin's negative feedback depth regulation model. The absolute distances moved before corrective vertical responses to threshold rates of pressure change are initated delimit the depth regulatory window. In darkness, the asymmetry of the window would lead to an ascent. It is hypothesized that light is an additional component in depth regulation, and that the limits and symmetry of the depth regulatory window may be controlled by the level of light adaptation.     

Occurrence of a shadow response among brachyuran larvae

A closed-circuit television system was used to study phototaxis and behavioral responses upon sudden decreases in light intensity in light-adapted Stage I zoeae from 7 species of Brachyura (Panopeus herbstii, Menippe mercenaria, Uca pugilator, Callinectes sapidus, Sesarma cinereum, Pinnotheres maculatum, and Libinia emarginata). All species except one show positive phototaxis to high intensities of 500 nm light and negative phototaxis to low intensities. Upon a sudden light intensity decrease, larvae show a shadow response, which depends upon the initial and final intensities. If the initial intensity is sufficient to induce positive phototaxis, and the light is extinguished, the larvae of 6 species stop swimming and passively sink (sinking response). Reducing the intensity to an absolute level that causes negative phototaxis, induces a sinking response followed by negative phototaxis. For intensity reductions that occur in the intensity range above that for negative phototaxis, a sinking responseis observed. The intensity decrease that initiates the sinking response is independent of initial stimulus intensity and duration, is consistent between species, and is equivalent to a decrease by 0.5 OD (optical density) units. The sinking response is greatest at intensity decreases of at least 1.0 to 1.1 OD units. It is concluded that the shadow response is common in brachyuran larvae inhabiting coastal-estuarine areas.     

Diel vertical migration of the marine copepod<Emphasis Type="Italic"> Calanopia americana</Emphasis>. II. Proximate role of exogenous light cues and endogenous rhythms

The marine copepod Calanopia americana Dahl undergoes twilight diel vertical migration (DVM) in the Newport River estuary, North Carolina, USA, in synchrony with the light:dark cycle. Copepods ascend to the surface at sunset, descend to the bottom around midnight, and make a second ascent and descent before sunrise. Behavioral assays with C. americana in the laboratory during fall 2002/2003 and summer 2004 investigated aspects of three hypotheses for the proximate role of light in DVM: (1) preferendum hypothesis (absolute irradiance), (2) rate of change hypothesis (relative rates of irradiance change), and (3) endogenous rhythm hypothesis. Results suggest that C. americana responds to exogenous light cues consistent with its DVM pattern; changes in absolute irradiance evoked swimming responses that would result in an ascent at sunset and descent at sunrise, while relative rates of irradiance decrease at sunset (–0.0046 s^–1) evoked an ascent response, and relative rates of irradiance increase at sunrise (0.0042 s^–1) evoked a descent response. Furthermore, C. americana expressed an endogenous rhythm in vertical migration that was positively correlated with field observations of twilight DVM. Collectively, these results indicate that both exogenous light cues and endogenous rhythms play a proximate role in twilight DVM of C. americana, providing redundancy in the causes of its vertical migration.Communicated by J.P. Grassle, New Brunswick     

The ontogeny of phototaxis by larvae of the crab Rhithropanopeus harrisii

Phototaxis by each zoeal stage of the crab Rhithropanopeus harrisii (Gould) was quantitatively measured by means of a microscope closed-circuit television system. The megalopa stage is indifferent to light stimulation and was thus not tested. The action spectrum for positive phototaxis is similar at each zoeal stage, having the most pronounced maximum at about 500 nm and smaller maxima at 400 and 280 nm. Responsiveness to various intensities of 500 nm light is also similar at each stage. After dark-adaptation strongest positive phototaxis occurs at intensities between 1.0 and 3.0x10^-4 W/m², with no negative phototaxis to lower intensities. After light-adaptation, the positive response occurs to higher intensities between 20 and 0.01 W/m², with a pronounced negative response to lower intensities. Generally, mean swimming speeds during positive phototaxis do not change with stimulation intensity, although during negative phototaxis mean speeds do vary with intensity. At the highest intensities which initiate a negative response, mean values are significantly greater than values for positive phototaxis. Based on the pattern of phototaxis at each developmental stage, a prediction of events during diurnal vertical migration is possible. The negative response after light-adaptation might also function as a shadow reflex.     

Depth regulation of larval marine decapod crustaceans: test of an hypothesis

This study tested the hypothesis that the dimensions and symmetry of the depth regulatory window of crustacean larvae are controlled by the level of light adaptation. Responses of first and last zoeal stages of the crab Rhithropanopeus harrisii (Gould) to different rates of pressure change were analyzed with a video system. Crabs were collected from the Neuse River estuary (North Carolina, USA) from May to September 1988. Responses were measured when larvae were adapted to light having an angular light distribution similar to that underwater at intensities ranging from one log unit above the lower phototaxis threshold to four log units higher. For both zoeal stages in darkness and at 10^-6 W m^-2, the distance larvae descend before responding to a pressure increase was much shorter than the distance they would ascend before responding to a pressure decrease. When adapted to a light level of 10^-4 W m^-2 both zoeal stages descended and ascended approximately equal distances before responding to an increase or decrease in pressure, respectively. Finally at the highest test light intensity (10^-2 W m^-2), the ascent distance was much shorter than the descent distance. These results support the hypothesis. The depth regulatory window dimensions predict an ascent in the water column upon adaptation to low light intensities and descent at high light levels. Thus Sulkin's negative feedback model provides the general mechanism of depth regulation. The effects of light adaptation on the limits of the depth regulatory window provide an additional component that negates the requirement for depth regulation at an absolute depth. The composite model can be termed the light-dependent negative feedback model of depth regulation.     

Copepod photobehavior in a simulated natural light environment and its relation to nocturnal vertical migration

To investigate the roles of light in initiating, controlling and directing nocturnal vertical migration, photoresponses of the adult, female copepod Acartia tonsa Dana were measured under simulated natural underwater light conditions using a video system. Copepods were adapted to a range of background light levels and tested with the following stimuli: absolute quantal intensity, absolute change in quantal intensity and relative (%) change in quantal intensity. The stimulus initiating vertical movements was relative change in quantal intensity, while responsiveness was controlled by the level of light adaptation. A. tonsa swam upward in response to decreases. Response with minimal stimulation occurred at an adaptation intensity close to that in the copepod's natural habitat at the time of the migratory ascent (near the bottom of the Newport River estuary, North Carolina, near sunset). Analysis of the angles of upward movement showed that light is not a directional cue. Relative increases in intensity resulted in sinking, with minimal stimulation required at an adaptation intensity close to that in the field when the migratory descent occurs near sunrise. These results offer a reasonable explanation of how light cues may shape nocturnal vertical migratory patterns.     

Effect of light intensity on activity and food-searching of larval herring,Clupea harengus: a laboratory study

Larvae of Clupea harengus were reared from spawning herring caught in March 1982 and 1983 in the Firth of Clyde, Scotland. An infra0red observation technique was used to record the behaviour of larval herring both in shallow dishes using a top view and in a tank 2 m deep using a side view. The amount of time larvae spent swimming, which was minimum in complete darkness, increased with increasing light intensity and as the larvae grew. Maximum swimming speeds of feeding larvae were recorded at light intensities between 10 and 100 lux. The presence of food organisms (Artemia sp., Brazilian strain) at light intensities below the feeding threshold (0.1 lux) caused an increase in the proportion of time spent active, but light intensities above the threshold had different effects, depending on developmental stage: larvae of 12 mm increased swimming speed, but 21 mm larvae decreased speed. In the 2 m deep tank in darkness, larvae displayed inactive periods wherein they sank head first, interspersed with periods of upward swimming. As light intensity increased, vertical swimming was replaced by horizontal swimming. These results are discussed with reference to food searching and vertical migration of larval herring in the sea.     

Responses of estuarine crab megalopae to pressure,salinity and light: Implications for flood-tide transport

The megalopal larval stage of many estuarine brachyuran crabs appears to return to adult habitats by undergoing rhythmic vertical migrations which result in saltatory up-estuary transport on flood tides. Larval ascent into the water column during rising tides may be cued by changing hydrologic variables. To test this hypothesis, we investigated the responses of field-caught megalopae of the blue crab Callinectes sapidus and the fiddler crab Uca spp. to constant rates of pressure and salinity change under laboratory conditions. For both genera, pressure changes resulted in increased movement (barokinesis) and upward migration in the test chamber, with C. sapidus megalopae having a lower response threshold (2.8×10^-2 mbar s^-1) than Uca spp. larvae (5×10^-2 mbar s^-1). Similarly, larvae ascended in response to increasing salinity, with C. sapidus larvae being more sensitive. Larvae were negatively phototactic and failed to respond to pressure increases at light levels above 1.0×10¹⁵ and 1.0×10¹³ photons m^-2 s^-1 for C. sapidus and Uca spp. megalopae, respectively. Such responses are thought to explain the low abundances of larvae in the water column during daytime flood tides. Nevertheless, threshold sensitivities to increasing pressure for both genera were above levels experienced during floodtide conditions in the field. Similarly, it is unlikely that increasing salinity is sufficient to induce ascent in Uca spp. postlarvae. However, rates of salinity increase during midflood tide typically reach levels necessary to induce an ascent in C. sapidus megalopae. These results are consistent with the hypothesis that fiddler crab megalopae utilize an endogenous activity rhythm for flood-tide transport, while blue crab megalopae rely upon external cues, especially salinity changes, to time their sojourns in the water column.     

Circatidal swimming behavior of brachyuran crab zoea larvae: implications for ebb-tide transport

Larvae of the blue crab Callinectes sapidus and fiddler crab Uca pugilator are exported from estuaries and develop on the continental shelf. Previous studies have shown that the zoea-1 larvae of some crab species use selective tidal-stream transport (STST) to migrate from estuaries to coastal areas. The STST behavior of newly hatched larvae is characterized by upward vertical migration during ebb tide followed by a descent toward the bottom during flood. The objectives of the study were (1) to determine if newly hatched zoeae of U. pugilator and C. sapidus possess endogenous tidal rhythms in vertical migration that could underlie STST, (2) to determine if the rhythms persist in the absence of estuarine chemical cues, and (3) to characterize the photoresponses of zoeae to assess the impact of light on swimming behavior and vertical distribution. Ovigerous crabs with late-stage embryos were collected from June to August 2002 and maintained under constant laboratory conditions. Following hatching, swimming activity of zoeae was monitored in darkness for 72 h. U. pugilator zoeae displayed a circatidal rhythm in swimming with peaks in activity occurring near the expected times of ebb currents in the field. Conversely, C. sapidus zoeae exhibited no clear rhythmic migration patterns. When placed in a light field that simulated the underwater angular light distribution, C. sapidus larvae displayed a weak positive phototaxis at the highest light levels tested, while U. pugilator zoeae were unresponsive. Swimming behaviors and photoresponses of both species were not significantly influenced by the presence of chemical cues associated with offshore or estuarine water. These results are consistent with predictions based on species-specific differences in spawning and the proximity of hatching areas to the mouths of estuaries. U. pugilator larvae are released within estuaries near the adult habitat. Thus, ebb-phased STST behavior by zoeae is adaptive since it enhances export. Selective pressures for a tidal migration in C. sapidus larvae are likely weaker than for U. pugilator since ovigerous females migrate seaward prior to spawning and hatching occurs near inlets and in coastal waters.     

Photosensitivity of the calanoid copepod Acartia tonsa

The light intensity and spectral sensitivities of the calanoid copepod Acartia tonsa Dana were determined by measuring phototactic responses. Adult females displayed only positive phototaxis. The dark-adapted copepod, which possesses a single naupliar eye, perceived light at intensities as low as 2.8x10¹¹ photons m^-2 s^-1. The action spectrum for positive phototaxis had no clear maxima but rather showed a broad range of greatest sensitivity from 453 to 620 nm. This sensitivity encompassed those wavelengths that are maximally available at the depth where the copepod is found during the day. This spectral overlap, coupled with the finding that the copepod requires light cues for nocturnal vertical miration, suggests that broad spectral sensitivity is an adaptive mechanism to maximize light intensity sensitivity during migration.     

Effects of the insect growth regulator Dimilin® (TH 6040) on larval development of two estuarine crabs

Effects of Dimilin^® (TH 6040), an insect growth regulator which interferes with the formation of the insect cuticle, were studied on the larval development of Rhithropanopeus harrisii (Gould) and Sesarma reticulatum (Say) (Crustacea: Brachyura). When larvae were exposed to 0.5 (R. harrisii only), 1, 3, 5, 7, and 10 ppb Dimilin from hatching to the first crab stage, survival in both species decreased in relation to increased concentrations of Dimilin. Survival of R. harrisii larvae wa significantly lower at 1 ppb and higher levels compared with control experiments, and in S. reticulatum a significant decrease in survival began at the 3 ppb level. At 10 ppb Dimilin, no larvae survived to the megalopa stage in either of the two species. The results indicate that early stage larvae of R. harrisii are more sensitive to Dimilin than those of S. reticulatum. When R. harrisii larvae were treated with 10 ppb Dimilin during the intermolt period of each of the 4 zoeal stages, nearly all larvae died during molting to the succeeding stage. First zoeal larvae of R. harrisii exposed to 10 ppb Dimilin at various days during the intermolt period were more sensitive to the compound late than early in the period. It is suggested that Dimilin also may interfere with the formation of the cuticle in crab larvae.     

Light responses of the larva of the serpulid polychaeteGaleolaria caespitosa

AdultGaleolaria caespitosa were collected at Cape Banks and Balmoral Bay, both in the vicinity of Sydney, Australia, from October 1985 to February 1986. Planktotrophic larvae of this serpulid polychaete are negatively phototactic at irradiance levels between 10 and 2 000x10¹⁴ q cm^-2 s^-1. The effective spectral range lies below 530 nm. Early metatrochophore larvae are less strongly photonegative; later they become photopositive. The photopositive phase persists into the actively swimming 3-setiger stage. Older setigerous larvae are photoneutral, demersal and positively thigmotactic. The strength of the photonegative response is not affected by the level of irradiance. The photopositive response is weaker at low levels of irradiance. The photonegative behaviour of young larvae may keep them at an optimal level for feeding. The subsequent photopositive phase may increase chances of being washed ashore and the terminal, photoneutral, thigmotactic phase may be important in the selection of a suitable site for settlement. Comparisons are made with light responses of two other serpulid species. Larval responses to light may be important infitness, especially in relation to successful niche occupation by adults.     

Responses of cirripede larvae to light. I. Experiments with white light

The phototactic response of the nauplius larva of Balanus balanoides, B. crenatus and Elminius modestus shows darkadaptation; the response of the cyprid of B. balanoides shows both phototaxis and low photokinesis. The phototactic responses and the orientation of the cyprid to white light at settlement require an intensity of illumination slightly above 10^-5 lux. The ability to select a shaded position by cyprids of R. balanoides requires a higher intensity of 10^-2 to 10^-4 lux; hence a different mechanism may be involved. Barnacle larvae are sufficiently sensitive to be able to respond to light beneath the sea surface, even on cloudy, moonless nights.     

Diel vertical migration of the marine copepod<Emphasis Type="Italic"> Calanopia americana</Emphasis>. I. Twilight DVM and its relationship to the diel light cycle

Marine copepods commonly exhibit vertical movements in the water column over the diel cycle, termed diel vertical migration (DVM), with the most common pattern being an ascent in the water column to minimum depth around sunset and descent to maximum depth around sunrise. The present study characterized the DVM pattern of the pontellid copepod Calanopia americana Dahl in the Newport River estuary (North Carolina, USA, in July 2003). The estuary is shallow and well-mixed, and the study site (34°43N; 76°40W), 1.5 km inside the estuary entrance, is unusual in lying within a gyre where tidal currents are always in the seaward direction. Changes in C. americana vertical abundance were related to spectrally relevant changes in light throughout the diel cycle. Simultaneous measurements of light and zooplankton abundance near the surface (0.5 m depth) and near the bottom (0.5 m above bottom) were made over one 4-h period and two 3-day periods during different phases of the tide. These observations suggest that C. americana undertook twilight DVM in the Newport River estuary; an ascent to the surface occurred at sunset, followed by a descent to near the bottom around midnight, with a second ascent to the surface and then descent to near bottom at sunrise. DVM in C. americana was independent of the tidal cycle, with the initial ascent in the water column at sunset possibly associated with relative rates of irradiance change. Copepod vertical movements were consistent with a night-active endogenous rhythm, and appeared independent of the abundance of predatory chaetognaths, Sagitta spp. In DVM studies with migrators like C. americana that are broadly sensitive to visible wavelengths of light, measuring photosynthetically active radiation may be a reasonable alternative to measuring light in a spectrally relevant photometric unit.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-005-1569-x.Communicated by J.P. Grassle, New Brunswick     

High-frequency observations of early-stage larval abundance: do storms trigger synchronous larval release in <Emphasis Type="Italic">Semibalanus balanoides</Emphasis>?

The acorn barnacle, Semibalanus balanoides, is thought to release larvae in response to phytoplankton blooms, but there is evidence that another, unidentified cue for release may exist. We conducted high-frequency sampling in Little Harbor, Massachusetts, USA, to determine whether early-stage larval abundance was related to several environmental variables, and to characterize vertical distributions of the larvae. Larval concentrations peaked at 2.52 and 1.02 individuals l⁻¹ during two storms. Larvae were more abundant near the surface than near the bottom. We suggest the hypothesis that turbid conditions and upward-swimming behavior may protect newly-released larvae from predation and cannibalism. Future studies should test this hypothesis with barnacles and other invertebrates.     

Comparative study of crustacean larval photoresponses

Ovigerous females of four brachyuran (Cancer gracilis, Lophopanopeus bellus bellus, Hemigrapsus oregonensis and Scyra acutiforns) and two anomuran (Pagurus beringanus and P. granosimanus) species, which live as adults in coastal areas, were collected near Friday Harbor, Washington, USA, in 1985, and spectral sensitivity and phototactic pattern of their larvae were measured. Responses were compared with previous measurements on estuarine species to determine whether responsiveness varies with adult habitat. Estuarine and coastal species have similar photoresponses. Spectral sensitivity of the test brachyran species had two maxima, one near 400 to 420 nm and another around 500 nm. The anomuran species were similar, but had an additional peak in the region of 580 to 620 nm. This sensitivity is adapted to daytime light conditions in the adult environment, and available spectra at the time of larval movement during diel vertical migration. Upon light adaptation and stimulation with a narrow light field, all six species showed positive phototaxis to high light intensities and a pronounced negative response to low intensities. Only the negative response will occur in natural underwater light conditions, and it is part of a predator-avoidance shadow-response which operates in areas of higher light intensity. The same phototactic pattern is observed in all species when darkadapted except H. oregonensis. If nocturnal vertical migration occurs, this negative response may be responsible for the descent at sunrise and depth maintenance during the day.     

Embryogenesis and larval biology of the ahermatypic scleractinian<Emphasis Type="Italic"> Oculina varicosa</Emphasis>

The ivory tree coral Oculina varicosa (Leseur, 1820) is an ahermatypic branching scleractinian that colonizes limestone ledges at depths of 6–100 m along the Atlantic coast of Florida. This paper describes the development of embryos and larvae from shallow-water O. varicosa, collected at 6–8 m depth in July 1999 off Fort Pierce, Florida (27°32.542 N; 79°58.732 W). The effect of temperature on embryogenesis, larval survival, and larval swimming speed were examined in the laboratory. Ontogenetic changes in geotaxis and phototaxis were also investigated. Embryos developed via spiral cleavage from small (100 µm), negatively buoyant eggs. Ciliated larvae developed after 6–9 h at 25°C. Embryogenesis ceased at 10°C, was inhibited at 17°C, and progressed normally at 25°C and 30°C. Larval survival, however, was high across the full range of experimental temperatures (11–31°C), although mortality increased in the warmest treatments (26°C and 31°C). Larval swimming speed was highest at 25°C, and lower at the temperature extremes (5°C and 35°C). An ontogenetic change in geotaxis was observed; newly ciliated larvae swam to the water surface and remained there for approximately 18 h, after which they swam briefly throughout the water column, then became demersal. Early larvae showed no response to light stimulation, but at 14 and 23 days larvae appeared to exhibit negatively phototactic behavior. Although low temperatures inhibited the development of O. varicosa embryos, the larvae survived temperature extremes for extended periods of time. Ontogenetic changes in larval behavior may ensure that competent larvae are close to the benthos to facilitate settlement. Previous experiments on survival, swimming speeds, and observations on behavior of O. varicosa larvae from deep-water adults indicate that there is no difference between larvae of the deep and shallow populations.Communicated by J.P. Grassle, New Brunswick     

Absence of sterol synthesis in larvae of the mud crab Rhithropanopeus harrisii and of the spider crab Libinia emarginata

A dietary requirement for sterols in crustacean larvae was established by the use of isotopic tracer techniques. Larvae of the mud crab Rhithropanopeus harrisii and larvae of the spider crab Libinia emarginata were exposed to acetate-¹⁴C or mevalonate-¹⁴C. Radioanalysis of the fatty acids of each species indicated sufficient incorporation of acetate for lipid synthesis. No radioactivity was detected in the sterols of animals exposed to either acetate-¹⁴C or mevalonate-¹⁴C. It was concluded that R. harrisii and L. emarginata larvae are unable to synthesize sterols from acetate or mevalonate and, therefore, require them in their diet.     

Ultrastructural study of the exoskeleton of the estuarine crab Rhithropanopeus harrisii: Effect of the insect growth regulator Dimilin® (diflubenzuron) on the formation of the larval cuticle

Ultrastructure of larval cuticle during the molt cycle of the estuarine crab Rhitropanopeus harrisii (Gould) (Crustacea: Brachyura) was studied in control larvae as well as in larvae exposed to 10 ppb of the insect growth regulator Dimilin^® (diflubenzuron). First zoeal larvae were used as test organisms. It has earlier been shown that 10 ppb Dimilin is lethal to zoeal larvae of R. harrisii, and nearly all exposed larvae died during molting to the next stage (Christiansen et al., 1978). Distinct differences in structure of the cuticle were found between the two groups of larvae. Both endocuticle and exocuticle appear to be deformed in Dimilin-treated larvae, whereas formation of epicuticle did not seem to be affected. The results indicate that Dimilin probably inhibits chitin synthesis in crab larvae as shown earlier by several authors for insect larvae.     

Behavioral response of fish larvae to low dissolved oxygen concentrations in a stratified water column

Density stratification and respiration lead to vertical gradients in dissolved oxygen in many aquatic habitats. The behavioral responses of fish larvae to low dissolved oxygen in a stratified water column were examined during 1990–1991 with the goal of understanding how vertical gradients in dissolved oxygen may directly affect the distribution and survival of fish larvae in Chesapeake Bay, USA. In addition, the effects of low oxygen on 24-h survival rates were tested so that results of behavior experiments could be interpreted in the context of risk to the larve. Naked goby [Gobiosoma bosc (Lacépède)] and bay anchovy [Anchoa mitchilli (Valenciennes)] larvae strongly avoided dissolved oxygen concentrations <1 mg 1^-1, which were lethal within 24 h at 25 to 27°C. In addition, naked goby larvae, whose behavior was tested at a wider range of dissolved oxygen concentrations, also showed a reduced preference for an oxygen concentration of 2 mg 1^-1, which leads to reduced survival during long-term exposures and to reduced feeding rates. There were no major differences in behavior or survival between the two species, or between the two age classes of naked gobies tested. Results suggest that behavioral responses to oxygen gradients will play a large role in producing marked vertical changes in abundance of feeding-stage larvae in Chesapeake Bay; mortality from direct exposure to low oxygen will likely be much less important in producing vertical patterns of larval abundance.