Depth regulatory behavior of the first stage zoea larvae of the sand crab Emerita analoga Stimpson (Decapoda: Hippidae)

Laboratory studies of the behavior of first stage zoea larvae of the sand crab Emerita analoga Stimpson have shown that while newly-hatched larvae are strongly photopositive, this response lasts only about four hours, as the larvae rapidly become photonegative. After becoming photonegative, a large proportion of the larvae remain so throughout the first four days of life if they are fed Artemia nauplii; if starved, the larvae become significantly more photopositive than when fed. Both the photopositive response of newly-hatched larvae and the reversal to photopositive behavior in response to starvation are only apparent under horizontal test conditions. Increases in hydrostatic pressure stimulate swimming activity among the larvae; responsiveness to pressure being greatest at hatching and decreasing thereafter. The pressure response is strongly oriented to light; pressure-stimulated larvae will swim towards a light source regardless of whether this involves upward, downward, or horizontal motion. Experiments suggest that the pressure response provides the primary mechanism for depth regulation among young larvae; gravity and light may augment the pressure ‘sense’ by serving as primary orientational cues. The nutritional status of an individual larva may alter its depth-regulatory capabilities, but this effect is not yet clear.     

Depth regulation of crab larvae in the absence of light

Behavioral responses to gravity and hydrostatic pressure have been investigated in two species of xanthid crabs Leptodius floridanus (Gibbes) and Panopeus herbstii Milne-Edwards to determine whether such responses provide a mechanism for depth regulation in the absence of light.In laboratory experiments, the four zoea stages and one megalopa stage of each species assume a differential vertical distribution in darkness, with succeeding stages showing a deeper overall distribution. Passive sinking rates increase in succeeding zoea stages and drop to an intermediate level after the molt to the megalopa stage. All zoea stages exhibit a negative geotaxis in the absence of light; the megalopa shows a positive geotaxis. The first zoea stage of Leptodius floridanus responds to an increase in hydrostatic pressure (up to 1 atmos above ambient) with an increase in swimming rate. This pressure response is shown to be reversible and not subject to short-term acclimation. The swimming rate of the last zoea stage does not increase in response to an increase in pressure.It is concluded that the responses of these larvae to gravity and hydrostatic pressure together with their characteristic passive sinking rates provide a mechanism for depth regulation in the absence of light that varies during ontogeny.     

Experimental predation studies of malacophagous larvae of Sepedon fuscipennis (Diptera: Sciomyzidae) and aquatic snails

Experimental studies were conducted under both laboratory and field conditions to determine the effects of prey density, three levels of prey aggregation, water depth, and predator density on the number of snails killed per larva of Sepedon fuscipennis. Of these factors, predation rates were most influenced by prey density and water depth. The number of small (2–4.5 mm) Lymnaea palustris killed per larva of S. fuscipennis increased at a decreasing rate as prey density increased under shallow water conditions. Larvae killed a mean of 14 snails at a prey density of 200/m², while an average of 24 snails were killed per larva of S. fuscipennis at a prey density of 4000/m². This functional response to prey density was largely confined to third-instar larvae, and as water depth was increased the response was not apparent.A field study in which larval densities of S. fuscipennis were manipulated showed that the population density of smaller individuals of L. palustris (< 4.5 mm) was reduced when predator density was increased. Populations of Physa integra, Gyraulus parvus, and larger L. palustris were not significantly reduced by the malacophagous larvae at the levels tested.     

Rapid behavioral responses of an invertebrate larva to dissolved settlement cue

Larvae of the nudibranch Phestilla sibogae were used to study whether a natural dissolved settlement cue (from their prey, Porites compressa, an abundant coral on Hawaiian reefs) induces behavioral responses that can affect larval transport to suitable settlement sites. As cue and larvae are mixed in the turbulent flow over a reef, cue is distributed in fine-scale filaments that the larva experiences as rapid (seconds) on/off encounters. To examine larval responses in this setting, individual larvae were tethered in a small flume with flow simulating water velocity relative to a freely swimming larva, and their responses to realistic temporal patterns of cue encounter were videotaped. Competent larvae quickly ceased swimming in cue filaments and resumed swimming after exiting filaments. The threshold cue concentration eliciting a response was 3%-17% of concentrations within heads of P. compressa in nature. When moving freely in filtered seawater, competent larvae swam along straight paths in all directions at approximately 0.2 cm s(-1), whereas in water conditioned by P. compressa, most ceased swimming and sank at approximately 0.1 cm s(-1). The ability of larvae to rapidly respond (by sinking) to brief encounters with dissolved settlement cues can enhance their rapid transport to the substratum, even in wave-driven turbulent flow.     

Behavioural significance of pressure responses in megalopa larvae of Callinectes sapidus and Macropipus sp.

The primarily benthic megalopa larvae of Callinectes sapidus and Macropipus holsatus swim upwards for a short distance from the bottom when subjected to pressure increased above ambient values, particularly when illuminated from above. Larvae exposed to stepwise increments of pressure exhibit a threshold in the barokinetic response which is lower in Callinectes (0.4 atm) than in Macropipus (0.8–1.0 atm). Such behaviour would tend to inhibit swimming in surface waters of stratified estuaries where net flow is seawards, but would promote slight upward swimming when on the bottom in deeper water which has a net flow inwards, thus substantiating other evidence that the megalopa is a stage at which recruitment to estuaries takes place. The lower threshold of the barokinetic response in Callinectes may be related to its occurrence in more uniformly shallow estuaries on the south and east coasts of N. America.

In both species swimming is markedly inhibited after a slight reduction in pressure. Such responses would be of value to larvae running the risk of being washed ashore in surf Sudden reductions in pressure by surf action would induce sinking into the undertow which would carry larvae offshore into deeper water.     

Temperature,salinity tolerance,and buoyancy during early development and starvation of Clyde and North Sea herring,cod, and flounder larvae

Tolerance limits, at which 50% of larvae could survive high temperature and low salinity for 24 h, were determined for the yolk-sac larvae of Clyde and North Sea herring (Clupea harengus L.), cod (Gadus morhua L.) and flounder (Platichthys flesus L.) during early development and starvation. Clyde and North Sea herring, cod and flounder from hatching to the end of the yolk-sac stage, could withstand 21–23.5 °C, 20.5–23 °C, 15.5–18 °C and 21.5–24°C, respectively. The temperature tolerance was reduced by about 3.5–4 °C for Clyde herring and cod, 4–4.5 °C for North Sea herring and 8–8.5 °C for flounder when the larvae reached the point-of-no-return (PNR, when 50% of larvae, although still alive, are no longer strong enough to feed). The lowest salinity tolerance between hatching and the end of yolk-sac stage was 1–1.5‰ for Clyde and North Sea herring, 2–3‰ for cod and 0–1‰ for flounder. In no instance was there a loss of tolerance to low salinity during starvation. In fact, tolerance improved somewhat until the larvae became moribund. At hatching Clyde and North Sea herring larvae were negatively buoyant with a sinking rate of 0.35–0.4cm · s⁻¹ which steadily decreased until the larvae became moribund. Cod and flounder larvae, however, were positively buoyant at hatching but became progressively less buoyant and, by the end of the yolk-sac stage they were negatively buoyant with a sinking rate of 0.06–0.07 cm · s⁻¹. This sinking rate then decreased slightly until the PNR stage. The low salinity tolerance of all three species varied in a similar fashion to buoyancy.     

Under pressure: biomechanical limitations of developing pneumatocysts in the bull kelp (Nereocystis luetkeana,Phaeophyceae)

Maintaining buoyancy with gas‐filled floats (pneumatocysts) is essential for some subtidal kelps to achieve an upright stature and compete for light . However, as these kelps grow up through the water column, pneumatocysts are exposed to substantial changes in hydrostatic pressure, which could cause complications as internal gases may expand or contract, potentially causing them to rupture, flood, and lose buoyancy. In this study, we investigate how pneumatocysts of Nereocystis luetkeana resist biomechanical stress and maintain buoyancy as they develop across a hydrostatic gradient. We measured internal pressure, material properties, and pneumatocyst geometry across a range of thallus sizes and collection depths to identify strategies used to resist pressure‐induced mechanical failure. Contrary to expectations, all pneumatocysts had internal pressures less than atmospheric pressure, ensuring that thalli are always exposed to a positive pressure gradient and compressional loads, indicating that they are more likely to buckle than rupture at all depths. Small pneumatocysts collected from depths between 1 and 9 m (inner radius = 0.4–1.0 cm) were demonstrated to have elevated wall stresses under high compressive loads and are at greatest risk of buckling. Although small kelps do not adjust pneumatocyst material properties or geometry to reduce wall stress as they grow, they are ~3.4 times stronger than they need to be to resist hydrostatic buckling. When tested, pneumatocysts buckled around 35 m depth, which agrees with previous measures of lower limits due to light attenuation, suggesting that hydrostatic pressure may also define the lower limit of Nereocystis in the field.     

Development of eggs of Antarctic krillEuphausia superba in relation to pressure

Summary This paper focuses upon the influence of increasing hydrostatic pressure on the development of krill eggs at 2°C. This experimental study on the embryology ofEuphausia superba was conducted at the Palmer Station, Antarctica during the 1982–1983 austral summer. The gravid females were captured from Bransfield Strait aboard theR/V Hero. The various embryological stages such as early cleavage, blastula, gastrula and limb-bud nauplius larva were defined and described. The duration for these various developmental stages at 1 atm was also established at +2°C and compared with the timing of this event at negative temperature. Krill embryonic development is inhibited at 4°C. The sinking rate of eggs and embryos was also measured at various pressure. The data suggest that pressure does not significantly influence the sinking rate. There appears to be a wide variation of sinking rates of eggs within the same brood. based on a simulated model of sinking rate, egg development was studied at increasing pressure. Pressure of 5–20 atm accelerates the rate of cleavage and therefore the 32-celled stage is attained within 5–8 h, while at 1 atm it took 13 h to reach the same stage. Pressure thus seems to have some influence on the duration of the development of different developmental stages of krill embryos.     

Effect of nutritional condition on larval food requisition behavior in a subterranean termite Reticulitermes speratus (Isoptera: Rhinotermitidae)

Although optimal investment theory would be similarly applicable to eusocial insects to maximize colony reproductive outputs, directly distinguishing an amount of investment in each larva should be a difficult task for workers because of the characteristics of group living. Thus, it is expected that workers adjust brood care by using a cue or signal conveying information of larval status. In termites, which are typical group of eusocial insects, there are nevertheless few direct observations on worker brood care and little is known about cues inducing worker feeding. I show here that a Japanese subterranean termite Reticulitermes speratus uses an overt food solicitation by larva, “pecking”, as a cue for worker feeding. Direct observations demonstrated that workers feed larvae in response to larval pecking. Furthermore, nutritional experiments showed that larvae exhibited pecking more frequently when their nutrient status is lower; hence, pecking may be an honest reflection of larval hunger status. These results indicate that workers can feed more starved larvae than less starved ones because pecking honestly reflects larval hunger state. That is, feeding in response to pecking should standardize the total amount of food intake of each larva and help a termite colony make worker investment efficient.     

Sinking rates of heterogeneous, temperate phytoplankton populations

Throughout the summer of 1978, the sinking rates of phytoplanktonwithin the Controlled Experimental Ecosystems (CEE's) were monitoredusing a technique based upon measurement of the transit timeof radioactively (¹⁴C) labeled cells. The experimental frameworkof FOODWEB 1 offered an unprecedented opportunity to documentthe sinking rates of heterogeneous phytoplankton of diversetaxonomic composition, growing under a variety of nutrient regimes;the absence of advective exchange in the CEE's provided knowledgeof the preconditioning history of the phytoplankton sampledat any given time. Sinking rates of whole phytoplankton assemblages (not size-fractioned)ranged from 0.32 – 1.69 m·day^–1; the averagerate (± s.d.) observed was 0.64 ± 0.31 m·day^–1.The most notable deviations from the mean value occurred whenthe population size distribution and taxonomic composition shifteddue to blooms. The relationship between phytoplankton sinkingand ambient nutrient levels was studied by following the ratesof a given size fraction (8–53 µm) for ten daysfollowing nutrient enrichment of a CEE. Over this time sinkingrates ranged from 1.08– 1.53 m·day^–1; decreasedrates occurred after nutrification, yet over the course of theentire trial sinking rates were not significantly (p >0.05)correlated to the ambient levels of any single nutrient species. The sinking rate implications of spore formation were also studied,and showed that sinking rates of Chaetoceros constrictus andC. socialis spores (2.75 ± 0.61 m·day^–1)were ca 5-fold greater than rates measured when the vegetativestages of these species dominated the population, reflectingthe influence of physiological mechanisms in controlling phytoplanktonbuoyancy. An example of the potential influence of colony formation uponbuoyancy was noted in observations of C. socialis which occasionallywas found to exist in large spherical configurations made ofcoiled cell chains and having low (0.40 m·day^–1)sinking rates. A hydrodynamic rationale is presented to showhow such a colony together with enveloped water may behave asa unit particle having lower excess density, and therefore lowobserved sinking rate, despite its conspicuously large size. Overall, sinking rates were not significantly correlated withany of the measured nutrient or photic parameters. There were,however, trials and comparisons which showed evidence for: (1)higher sinking rates in populations dominated by large cells,(2) decreased sinking rates after nutrient enrichment, and (3)buoyancy response to light levels. It is suggested that correlationof sinking rates with synoptic environmental measurements atany given time is not explicit because the associations mayinvoke lag times of physiological response. The interpretationmade from these findings is that the preconditioning historyof the population, rather than the prevailing conditions atthe time of a given measurement, is most closely associatedwith population buoyancy modifications.     

Juvenile hormone and the induction of larval polymorphism and the diapause of the southwestern corn borer, Diatraea grandiosella

The southwestern corn borer, Diatraea grandiosella, enters diapause as an immaculate mature larva which is a polymorphic variant of the spotted non-diapause larva. Because dormant immaculate larvae could be obtained by treating last stage non-diapause larvae with a juvenile hormone mimic (JHM), experiments were conducted to determine whether these hormonally induced immaculate larvae (HIL) were physiologically comparable to the normal environmentally induced immaculate diapause larvae (EIL). Comparative data obtained about pupation rates, response to JHM, metabolic reserves, oxygen consumption, and the state of spermatogenesis of HIL and EIL led to the conclusion that the HIL were physiologically similar to the EIL. The results demonstrate that the developmental programme of non-diapause larvae could be switched and ‘diapause’ induced solely by the topical application of JHM. We believe that the data further support our hypothesis that the larval diapause of D. grandiosella is initiated and maintained by the juvenile hormone.     

Steady-state pleural fluid flow and pressure and the effects of lung buoyancy

Both theoretical and experimental studies of pleural fluid dynamics and lung buoyancy during steady-state, apneic conditions are presented. The theory shows that steady-state, top-to-bottom pleural-liquid flow creates a pressure distribution that opposes lung buoyancy. These two forces may balance, permitting dynamic lung floating, but when they do not, pleural-pleural contact is required. The animal experiments examine pleural-liquid pressure distributions in response to simulated reduced gravity, achieved by lung inflation with perfluorocarbon liquid as compared to air. The resulting decrease in lung buoyancy modifies the force balance in the pleural fluid, which is reflected in its vertical pressure gradient. The data and model show that the decrease in buoyancy with perfluorocarbon inflation causes the vertical pressure gradient to approach hydrostatic. In the microgravity analogue, the pleural pressures would be toward a more uniform distribution, consistent with ventilation studies during space flight. The pleural liquid turnover predicted by the model is computed and found to be comparable to experimental values from the literature. The model provides the flow field, which can be used to develop a full transport theory for molecular and cellular constituents that are found in pleural fluid.     

EFFECTS OF ENVIRONMENTAL VARIATION ON SINKING RATES OF MARINE PHYTOPLANKTON1

The effects of environmental variables, particularly irradiance, on the sinking rates of phytoplankton were investigated using cultures of Chaetoceros gracilis Schütt and C. flexuosum Mangin in laboratory experiments; these data were compared with results from assemblages in the open ocean and marginal ice zone of the Greenland Sea. In culture experiments both the irradiance under which the diatom was grown and culture growth rate were positively correlated with sinking rates. Sinking rates (ψ) in the Greenland Sea were smallest when determined from chlorophyll (mean ψ_chl= 0.14 m · d^?1) and biogenic silica (ψ_si= 0.14 m · d^?1) and greatest when determined from particulate carbon (ψ_c= 0.55 m · d^?1) and nitrogen (ψ_N= 0.64 m · d^?1). Field measurements indicated that variations in sinking may be associated with changes in irradiance and nitrate concentrations. Because these factors do not directly affect water density, they must be inducing physiological changes in the cell which affect buoyancy. Although a direct response to a single environmental variable was not always evident, sinking rates were positively correlated with growth rates in the marginal ice zone, further indicating a connection to physiological processes. Estimats of carbon flux at stations with vertically mixed euphotic zones indicated that approximately 30% of the daily primary production sank from the euphotic zone in the form of small particulates. Calculated carbon flux tended to increase with primary productivity.     

Entrainment of the larval release rhythm of the crab Rhithropanopeus harrisii (Brachyura: Xanthidae) by cycles in hydrostatic pressure

This study investigated the entrainment of a larval release rhythm by determining whether a tidal cycle in hydrostatic pressure could entrain the circatidal rhythm in larval release by the crab Rhithropanopeus harrisii (Gould). Ovigerous females were collected from a non-tidal estuary. The time of larval release by individual crabs was monitored under constant conditions with a time-lapse video system. Crabs with mature embryos at the time of collection had a pronounced circadian rhythm in larval release with a free running period of 25.1 h. Crabs with immature embryos that were maintained under constant conditions from the time of collection until larval release retained a weak circadian rhythm. Other crabs with immature embryos were exposed to a tidal cycle in step changes in hydrostatic pressure equivalent to 1 m of water. This cycle entrained a circatidal rhythm in larval release. The free-running period was 12.1 h and larvae were released at the time of the transition from low to high pressure. Although past studies demonstrated that a tidal cycle in hydrostatic pressure could entrain activity rhythms in crustaceans, this is the first study to show that pressure can entrain a larval release rhythm.     

Effects of pressure on swimming behavior in planula larvae of the coral Porites astreoides (Cnidaria, Scleractinia)

Mechanisms governing the behavior of coral planulae are not well understood, particularly those manifesting themselves between the time when the larvae are released and when they settle. Larvae from the hermatypic coral Porites astreoides Lamarck were exposed to different levels of hydrostatic pressure—103.4, 206.9, 310.3, 413.8, and 517.1 kPa (including ambient pressure). Data were collected at stops of the above pressures for 15 min each, respectively. This was done in both an increasing sequence and a decreasing one. When exposed to increases in pressure from 103.4 kPa, larvae swam upward (negative barotaxis) in a spiraling motion. Upon exposure to decreasing pressure from 517.1 kPa, larvae moved downward (positive barotaxis), but the magnitude of the vertical movement was much less than in the case of increasing pressure. This suggests that these larvae are more sensitive to increased pressure than decreasing pressure. High variance was also observed in the responses of these larvae at both the intra- and inter-colony levels. Thus, this behavioral trait is variable within the population. The trait may be genetically based, and thus may be susceptible to alteration by natural selection, although this remains to be demonstrated. This study is the first to document these behavioral mechanisms in coral larvae.     

Planktothrix populations in subalpine lakes: selection for strains with strong gas vesicles as a function of lake depth,morphometry and circulation

1. The genus Planktothrix (Cyanobacteria) usually produces concentrated populations of filaments in the summer metalimnion of thermally stratifying lakes. This has been associated with the action of gas vesicles, cellular structures providing positive buoyancy. At the end of the summer, filaments are carried by convective mixing deeper into the water column where some gas vesicles collapse as a result of high hydrostatic pressure. They then lose their buoyancy, sink and are lost from the euphotic zone. 2. The resistance of gas vesicles to hydrostatic pressures is critical for the survival of Planktothrix in deep lakes. However, comparative observations on populations from lakes of a range of depths and hydrodynamic regimes are still needed to examine the relationships between the adaptive trait (i.e. the ‘critical’ pressure at which each gas vesicle collapses) with the environmental factor (i.e. the maximum hydrostatic pressure). 3. To explore the adaptation of Planktothrix populations to the depth of winter circulation in different systems, we collected 276 strains of P. cf. rubescens from eight lakes (z_max = 24–410 m) in Northern Italy during summer 2009 and we analysed the multicopy gene gvpC coding for a protein that crucially influences the critical pressure. 4. The strains analysed clustered into two main groups having gas vesicles with a mean critical pressure of 1.1 and 0.9 MPa, respectively. The proportion of the stronger strains was generally positively related to lake depth, although the overall pattern was complicated by individual lake morphology and hydrology. The relative frequency of stronger filaments was (i) greatest in deep basins with concave slopes and (ii) least in one deep, but permanently stratified lake. 5. The simultaneous presence of ‘weaker’ and ‘stronger’ filaments could allow for a rapid adaptive response to changes in hydrostatic pressures, related to changes in the amplitude of vertical circulation characterising deep lakes.     

Synchronous Effects of Temperature, Hydrostatic Pressure, and Salinity on Growth, Phospholipid Profiles, and Protein Patterns of Four Halomonas Species Isolated from Deep-Sea Hydrothermal- Vent and Sea Surface Environments

Four strains of euryhaline bacteria belonging to the genus Halomonas were tested for their response to a range of temperatures (2, 13, and 30°C), hydrostatic pressures (0.1, 7.5, 15, 25, 35, 45, and 55 MPa), and salinities (4, 11, and 17% total salts). The isolates were psychrotolerant, halophilic to moderately halophilic, and piezotolerant, growing fastest at 30°C, 0.1 MPa, and 4% total salts. Little or no growth occurred at the highest hydrostatic pressures tested, an effect that was more pronounced with decreasing temperatures. Growth curves suggested that the Halomonas strains tested would grow well in cool to warm hydrothermal-vent and associated subseafloor habitats, but poorly or not at all under cold deep-sea conditions. The intermediate salinity tested enhanced growth under certain high-hydrostatic-pressure and low-temperature conditions, highlighting a synergistic effect on growth for these combined stresses. Phospholipid profiles obtained at 30°C indicated that hydrostatic pressure exerted the dominant control on the degree of lipid saturation, although elevated salinity slightly mitigated the increased degree of lipid unsaturation caused by increased hydrostatic pressure. Profiles of cytosolic and membrane proteins of Halomonas axialensis and H. hydrothermalis performed at 30°C under various salinities and hydrostatic pressure conditions indicated several hydrostatic pressure and salinity effects, including proteins whose expression was induced by either an elevated salinity or hydrostatic pressure, but not by a combination of the two. The interplay between salinity and hydrostatic pressure on microbial growth and physiology suggests that adaptations to hydrostatic pressure and possibly other stresses may partially explain the euryhaline phenotype of members of the genus Halomonas living in deep-sea environments.     

An offensive predator phenotype selects for an amplified defensive phenotype in its prey

Inducible defenses of prey and inducible offenses of predators are examples of adaptive phenotypic plasticity. Although evolutionary ecologists have paid considerable attention to the adaptive significances of these strategies, they have rarely focused on their evolutionary impacts on the interacting species. Because the functional phenotypes of predator and prey determine strength of interactions between the species, the inducible plasticity can modify selective pressure on trait distribution and, ultimately, trait evolution in the interacting species. We experimentally tested this hypothesis in a predator–prey system composed of salamander larvae (Hynobius retardatus) and frog tadpoles (Rana pirica) capable of expressing antagonistic inducible offensive or defensive traits, an enlarged gape in the salamander larvae and a bulgy body in the tadpoles, when predator–prey interactions are strong. We examined selection strength on the tadpole’s defensive trait by comparing survival rates of tadpoles with different defensive levels under predation pressure from offensive or non-offensive salamander larvae. Survival rates of more-defensive tadpoles were greater than those of less-defensive tadpoles only when the tadpoles were exposed to offensive salamander larvae; thus, the predator’s offensive phenotype could select for an amplified defensive phenotype in their prey. As the expression of inducible offenses by H. retardatus larvae depends greatly on the composition of its ecological community, the inducible defensive bulgy morph of R. pirica tadpoles might have evolved in response to the variable expression of the H. retardatus offensive larval phenotype.     

Effects of water temperature on cardiac autonomic nervous system modulation during supine floating

It is well known that heart rate, oxygen uptake and body temperature during exercise in water are affected by water temperature, buoyancy, hydrostatic and so on. It has been reported that the central blood volume during immersion was affected by the increased external hydrostatic pressure and cold-induced peripheral vasoconstriction, and intrathoratic blood volume should be greater during cold than warm water immersion (Epstein, 1992). The purpose of this presentation study was to make clear heart rate, blood pressure, oxygen uptake and cardiac autonomic nervous system modulation during supine floating at water temperatures of 25, 35 and 41 degrees C.     

Osmotic Behavior of Oat Coleoptile Tissue in Relation to Growth

Efforts were made to estimate the water potential difference that is required, between rapidly growing oat coleoptile cylinders and dilute medium, to support the rate of water uptake involved in elongation, (a) by the traditional method of determining the concentration of mannitol in which the tissue neither gains nor loses water, and (b) by measuring the rates of osmotic exchanges induced by treating the tissue with different hypotonic mannitol concentrations. Both methods indicated large water potential differences (3 to 10 atm), in some cases approaching the osmotic pressure of the cells. However, indication was obtained that the rates of osmotic exchanges induced by mannitol solutions, and presumably also the equilibrium response sought in (a), are governed by the rate of diffusional exchange of mannitol with the free space rather than by the permeability of the tissue to water. Osmotic swelling of the tissue measured by immersing it in water after its turgor pressure had been reduced by evaporation, was at least two to four times more rapid than when mannitol was involved. The permeability to water estimated by the evaporation-immersion method indicated that rapidly elongating cylinders have water potentials between -0.8 and -2.5 atm, or between 10 and 25 per cent of their osmotic pressure.