SENSORY EQUILIBRIUM AND DARK ADAPTATION IN MYA ARENARIA

1. The reaction time of Mya to light is composed of two parts. The first, a sensitization period, is an exceedingly short interval of the order of magnitude associated with photographic processes. The second is a latent period of about 1.3 seconds, during which Mya need not remain exposed to the stimulating light. 2. The process of dark adaptation in Mya is orderly. Its progress may be represented by the formation of a photosensitive substance according to the dynamics of a bimolecular reaction. See PDF for Structure 3. Photosensory equilibrium as represented by the light- and dark-adapted conditions finds a rational explanation in terms of the "stationary state" of a reversible photochemical reaction involving a photosensitive substance and its two precursors. 4. There are two corollaries to this hypothesis. The first requires that the reaction time at sensory equilibrium for a given intensity should vary inversely with the temperature; the second, that the rate of dark adaptation should vary directly with the temperature. Experiments verified both of these requirements.     

THE EFFECT OF EXPOSURE PERIOD AND TEMPERATURE ON THE PHOTOSENSORY PROCESS IN CIONA

1. Experiments are presented which show that the latent period in the photosensory response of Ciona is inversely proportional to the duration of the exposure period to light. From this it is found that the velocity of the chemical reaction which determines the latent period is directly proportional to the concentration of photochemical products formed during the exposure period. This is interpreted as showing that the two processes form a coupled photochemical reaction, of which the secondary reaction proceeds only in the presence of products from the primary reaction. This coupling may be a catalysis or a direct chemical relation. 2. Further experiments show that the relation between temperature and the latent period is accurately described by the Arrhenius equation in which µ = 16,200. The precise numerical value of µ tentatively identifies the latent period process as an oxidation reaction which is catalyzed by iron. 3. The photocatalytic properties of certain iron compounds are used as a model for the coupled photochemical reaction suggested for the photosensory mechanism of Ciona and Mya.     

DARK ADAPTATION AND THE LIGHT-GROWTH RESPONSE OF PHYCOMYCES

1. A single-celled, elongating sporangiophore of Phycomyces responds to a sufficient increase in intensity of illumination by a brief increase in growth rate. This is the "light-growth response" of Blaauw. 2. The reaction time is compound, consisting of an exposure period and a latent period (this comprising both the true latent period resulting from photochemical action and any "action time" necessary for the response). During the latter period the plant may be in darkness, responding nevertheless at the end of the latent period. 3. Both light adaptation and dark adaptation occur in the sporangiophore. The kinetics of dark adaptation can be accounted for on the basis of a bimolecular reaction, perhaps modified by autocatalysis. Attention is called to the bimolecular nature of the "dark" reaction in all other photosensory systems that have been studied, in spite of the diversity of the photosensitive substances themselves and of the different forms of the responses to light.     

DARK ADAPTATION AND THE DARK GROWTH RESPONSE OF PHYCOMYCES

Light-adapted sporangiophores of the fungus Phycomyces respond to sudden darkening by a temporary decrease in the rate of elongation, after a latent period of several minutes. The reaction time of this "dark growth" response is compound like that of the "light growth" response. It is, moreover, shorter the more intense the previous illumination. The rate of dark adaptation following adaptation to a very large range of light intensities is found to be proportional to the logarithm of the preceding light intensity. It is shown that a constant amount of dark adaptation takes place before the response occurs. On the assumption that changes in the rate of growth reflect changes in the concentration of a substance which at constant light intensity is in equilibrium with a light-sensitive material, possible equations for such a photostationary state are examined. The most reasonable formulation requires that the partial velocity of the "light" reaction be taken proportional to log I instead of to I directly.     

Sensitivity to light in the sea anemone Metridium senile (L). II. Studies of reaction time variability and the effects of changes in light intensity and temperature

1. The reaction time of the photosensitive response of Metridium was found to be composed almost entirely of a sensitization period. If a latent period exists, it is too short to be detected by the methods used and is probably less than a second in duration. The length of the reaction time, therefore, was used as a measure of the radiant energy necessary at any intensity to elicit a response. 2. The length of the reaction time was found to vary randomly by a factor of seven under constant environmental and stimulating conditions. Determination of large numbers of reaction times on several anemones, grouped according to length, gave closely similar distributions resembling Poisson distributions. It was suggested that the variability may be caused by the same factor or mechanism in each individual. An experimental scheme was presented for determining the expected error and variation in statistical quantities when groups of ten observations are used. 3. The means of groups of reaction times determined at different intensities formed a hyperbolic relationship when plotted against intensity, suggesting that the animal obeys the Bunsen-Roscoe law of reciprocity. No marked changes were noted of per cent deviation from the mean or of randomness at different intensities. Stimulation of Metridium requires roughly 5 x 10⁹ incident quanta/cm.² of blue-green light. 4. No temperature effect could be found on either the means, the per cent deviation from the mean, or the degree of randomness of series of about ten observations, studied over a 17°C. range. It was concluded that the variability in reaction time was not due to changes in any complex physiological state such as muscular tone. 5. A possible use of the photosensitive response is suggested and the potentialities of "integrative" photosensitivity are discussed. 6. Possible mechanisms to explain the variability in reaction time are discussed. In the light of evidence presented, the most likely hypothesis appears to be the uncertainty of quantum capture caused by low concentrations of photosensitive pigment. Assuming the validity of this hypothesis, evidence suggests that the anemones responded to less than 10 quanta of absorbed light. Caution, however, is recommended in accepting the hypothesis because of the indirect nature of the supporting evidence.     

THE INFLUENCE OF TEMPERATURE ON THE PHOTOSENSORY LATENT PERIOD

1. The effect of temperature on the photosensory latent period in Pholas dactylus is accurately described by the Arrhenius equation when µ = 18,300. 2. The adequacy of this equation has already been found for two other photosensitive animals, Mya and Ciona, which are very similar in behavior to Pholas. The value of µ is different for each of the three species studied. 3. This is taken to mean that though the organization of the receptor process is the same for the three species, the chemical materials concerned are very likely different.     

THE KINETICS OF DARK ADAPTATION

1. Data are presented for the dark adaptation of four species of animals. They show that during dark adaptation the reaction time of an animal to light of constant intensity decreases at first rapidly, then slowly, until it reaches a constant minimum. 2. On the assumption that at all stages of adaptation a given response to light involves a constant photochemical effect, it is possible to describe the progress of dark adaptation by the equation of a bimolecular reaction. This supposes, therefore, that dark adaptation represents the accumulation within the sense cells of a photosensitive material formed by the chemical combination of two other substances. 3. The chemical nature of the process is further borne out by the fact that the speed of dark adaptation is affected by the temperature. The velocity constant of the bimolecular process describing dark adaptation bears in Mya a relation to the temperature such that the Arrhenius equation expresses it with considerable exactness when µ = 17,400. 4. A chemical mechanism is suggested which can account not only for the data of dark adaptation here presented, but for many other properties of the photosensory process which have already been investigated in these animals. This assumes the existence of a coupled photochemical reaction of which the secondary, "dark" reaction is catalyzed by the products of the primary photochemical reaction proper. This primary photochemical reaction itself is reversible in that its main products combine to form again the photosensitive material, whose concentration controls the behavior of the system during dark adaptation.     

THE DARK ADAPTATION OF THE HUMAN EYE

During the dark adaptation of the human eye, its visual threshold decreases to a small fraction of its original value in the light. An analysis of the quantitative data describing this adaptation shows that it follows the course of a bimolecular chemical reaction. On the basis of these findings it is suggested that visual reception in dim light is conditioned by a reversible photochemical reaction involving a photosensitive substance and its two products of decomposition. Accordingly, dark adaptation depends on the course of the "dark" reaction during which the two products of decomposition reunite to synthesize the original photosensitive substance.     

THE EFFECT OF TEMPERATURE ON THE LATENT PERIOD IN THE PHOTIC RESPONSE OF MYA ARENARIA

1. The effect of temperature on the reaction time of Mya to light is mainly confined to the latent period. The sensitization period, representing a photochemical process, is changed comparatively little. 2. The relation between the latent period and the temperature is adequately expressed by the Arrhenius equation, for temperatures below 21°C. Above this temperature, the latent period becomes increasingly longer than is required by the Arrhenius formula when µ = 19,680. 3. These deviations, occurring above the highest environmental temperature of Mya, are explained on the assumption that the principal product formed during the latent period is inactivated by heat. 4. Calculation of the velocity of the hypothetical inactivation reaction at different temperatures shows that it also follows the Arrhenius rule when µ = 48,500. This value of µ corresponds to those generally found for spontaneous inactivations and destructions.     

Characteristics of intracellular on - off responses of amacrine cells of the dark-adapted carp retina

The dependence of the amplitude and latent period of intracellular on and off responses of the amacrine cells of the isolated, dark-adapted carp retina on the intensity and diameter of the light spot was investigated. On and off responses of amacrine cells to light were shown to consist of fast depolarization responses with oscillations and spikes superposed upon them. With an increase in the intensity and area of the stimulus the latent period of the on response decreases but that of the off response increases. A near-linear relationship was found between the amplitude of the on response and the logarithm of the diameter of the spot up to 3 mm during changes in stimulus intensity of not more than 4 logarithmic units. With an increase in stimulus intensity the amplitude and zone of summation of the off response are reduced; it is suggested that under these circumstances this decrease may be connected with the different amplitude and temporal characteristics of off processes in the bipolar cells converging on the amacrine cells.     

A LATENT PERIOD IN THE ACTION OF COPPER ON RESPIRATION

1. When copper chloride is allowed to act on Aspergillus niger there is at first a period during which there is no change in the rate of the production of carbon dioxide, following which the rate of respiration falls. The interval of no change is called the latent period. 2. When the copper is removed from the external solution before the end of the latent period this interval is prolonged. The rate of respiration then falls to a new level below the normal level. 3. Experiments on Nitella and on Valonia indicate that the copper penetrates the cell almost immediately. 4. The length of the latent period varies inversely as a constant power of the concentration of the copper. 5. These results are explained by assuming that the copper is made active in the respiration system by means of a reversible reaction. By using appropriate velocity constants the experimental curves can be duplicated by calculated curves.     

Fluorescent Protein Voltage Probes Derived from ArcLight that Respond to Membrane Voltage Changes with Fast Kinetics

We previously reported the discovery of a fluorescent protein voltage probe, ArcLight, and its derivatives that exhibit large changes in fluorescence intensity in response to changes of plasma membrane voltage. ArcLight allows the reliable detection of single action potentials and sub-threshold activities in individual neurons and dendrites. The response kinetics of ArcLight (τ1-on ~10 ms, τ2-on ~ 50 ms) are comparable with most published genetically-encoded voltage probes. However, probes using voltage-sensing domains other than that from the Ciona intestinalis voltage sensitive phosphatase exhibit faster kinetics. Here we report new versions of ArcLight, in which the Ciona voltage-sensing domain was replaced with those from chicken, zebrafish, frog, mouse or human. We found that the chicken and zebrafish-based ArcLight exhibit faster kinetics, with a time constant (τ) less than 6ms for a 100 mV depolarization. Although the response amplitude of these two probes (8-9%) is not as large as the Ciona-based ArcLight (~35%), they are better at reporting action potentials from cultured neurons at higher frequency. In contrast, probes based on frog, mouse and human voltage sensing domains were either slower than the Ciona-based ArcLight or had very small signals.     

THE NATURE OF FOVEAL DARK ADAPTATION

1. After a discussion of the sources of error involved in the study of dark adaptation, an apparatus and a procedure are described which avoid these errors. The method includes a control of the initial light adaptation, a record of the exact beginning of dark adaptation, and an accurate means of measuring the threshold of the fovea after different intervals in the dark. 2. The results show that dark adaptation of the eye as measured by foveal vision proceeds at a very precipitous rate during the first few seconds, that most of the adaptation takes place during the first 30 seconds, and that the process practically ceases after 10 minutes. These findings explain much of the irregularity of the older data. 3. The changes which correspond to those in the fovea alone are secured by correcting the above results in terms of the movements of the pupil during dark adaptation. 4. On the assumption that the photochemical effect of the light is a linear function of the intensity, it is shown that the dark adaptation of the fovea itself follows the course of a bimolecular reaction. This is interpreted to mean that there are two photolytic products in the fovea; that they are disappearing because they are recombining to form anew the photosensitive substance of the fovea; and that the concentration of these products of photolysis in the sense cell must be increased by a definite fraction in order to produce a visual effect. 5. It is then suggested that the basis of the initial event in foveal light perception is some mechanism that involves a reversible photochemical reaction of which the "dark" reaction is bimolecular. Dark adaptation follows the "dark" reaction; sensory equilibrium is represented by the stationary state; and light adaptation by the shifting of the stationary state to a fresh point of equilibrium toward the "dark" side of the reaction.     

GEOTROPIC EXCITATION IN HELIX

Rotation of an inclined surface on which Helix is creeping straight upward, such that the axis of the animal is turned at a right angle to its previous position, but in the same plane, leads to negatively geotropic orientation after a measurable latent period or reaction time. The duration of the latent period is a function of the slope of the surface. The magnitude of the standard deviation of the mean latent period is directly proportional to the mean latent period itself, so that the relative variability of response is constant. The dependence of reaction time upon extent of displacement from symmetrical orientation in the gravitational field is found also by tilting the supporting surface, without rotation in the animal''s own plane. On slopes up to 55°, the relation between latent period and the sine of the slope is hyperbolic; above this inclination, the latent period sharply declines. This change in the curve is not affected by the attachment of moderate loads to the snail''s shell (up to 1/3 of its own mass), and is probably a consequence of loss of passive stable equilibrium when rotated. When added loads do not too greatly extend the snail''s anterior musculature, the latent period for the geotropic reaction is decreased, and, proportionately, its σ. These facts are discussed from the standpoint that geotropic excitation in these gasteropods is due to impressed muscle-tensions.     

Effect of photoperiod on a winter and on a summer diapause in two species of cranefly (Tipulidae)

Two craneflies, Tipula subnodicornis and Tipula pagana, both undergo diapause in the final larval instar. The species showed differences in the intensity of diapause and in the timing of the photoperiodic reaction during diapause, that could be related to season. Tipula subnodicornis undergoes a winter diapause that is induced and maintained in its early stages by short photoperiod (L:D:6:18). In the laboratory individuals in the early stages of diapause terminated diapause and pupated earlier when they were exposed to daylengths of, or greater than, 12 hr. However, it is suggested that in the field diapause is broken before the natural daylength is long enough to have any accelerating effect on development. Tipula pagana has a summer diapause which is of greater intensity than that of Tipula subnodicornis and some larvae were maintained for 197 days in the laboratory, without pupating, on an 18 hr daylength. Diapause was broken by a L:D;16:8 photoperiod and development was accelerated by a further decrease in daylength. The acceleration in development rate was attended by a decrease in the variance about the mean date of emergence and resulted in a highly synchronised emergence period. It is suggested that this quantitative response to daylength is particularly important to a species that emerges in the autumn when the temperature in the field is falling.     

The latent period of anode break excitation in myelinated and giant axons

The time lapse (latent period) between a hyperpolarizing pulse and anodic break response was investigated in single myelinated (frog) and giant axons (squid). The latent period was found to first decrease when the stimulating current intensity was gradually increased beyond threshold. However, for higher stimulus intensities the latent period increased and became a direct function of stimulus intensity. These findings were shown to be consistent with the Hodgkin-Huxley axon model.The initial decrease in latency was shown to be related to the changes in potassium current. The following increase in latency was due to the passive cable properties of the fibres. The increased tendency of sensory and other fibers to fire repetitively is analysed in view of the above findings.     

Predator overcomes the Allee effect due to indirect prey–taxis

A mathematical model for spatiotemporal dynamics of prey–predator system was studied by means of linear analysis and numerical simulations. The model is a system of PDEs of taxis–diffusion–reaction type, accounting for the ability of predators to detect the locations of higher prey density, which is formalized as indirect prey–taxis, according to hypothesis that the taxis stimulus is a substance being continuously emitted by the prey, diffusing in space and decaying with constant rate in time (e.g. odour, pheromone, exometabolit). The local interactions of the prey and predators are described by the classical Rosenzweig – MacArthur system, which is modified in order to take into account the Allee effect in the predator population. The boundary conditions determine the absence of fluxes of population densities and stimulus concentration through the habitat boundaries. The obtained results suggest that the prey–taxis activity of the predator can destabilize both the stationary and periodic spatially-homogeneous regimes of the species coexistence, causing emergence of various heterogeneous patterns. In particular, it is demonstrated that formation of local dense aggregations induced by prey–taxis allows the predators to overcome the Allee effect in its population growth, avoiding the extinction that occurs in the model in the absence of spatial effects.     

The dynamics of latent inhibition in rats under the action of substance P]

The role of substance P in latent inhibition was studied in experiments on rats. Administration of neuropeptide during pre-exposition of conditioned stimulus and before conditioning disturbed all signs of latent inhibition: level of reproduction, retention and resistance to amnestic action of conditioned reaction in the task of passive avoidance. Single administration of haloperidol before learning prevented the disturbance. Significance of hyperfunction of substance P in selective attention and pathogenesis of schizophrenia is discussed.     

THE LIGHT GROWTH RESPONSE AND THE GROWTH SYSTEM OF PHYCOMYCES

1. The Roscoe-Bunsen law holds for the light growth response of Phycomyces if the time component of stimulation is short. With exposures longer than a few seconds, the reaction time to light is determined by the intensity and not by the energy of the flash. 2. The possible nature of the very long latency in the response to light is considered in terms of the structure of the cell and its mechanism of growth. It is suggested that during the latency some substance produced by light in the protoplasm is transported centrifugally to the cell wall or outermost layer of protoplasm. 3. The total elongation occurring over a period of 1 to 2 hours is independent of flashes of light or temporary darkening. Light acts by facilitating some change already under way in the growth system, and during the principal phase of elongation is not a necessary or limiting factor for growth. 4. Judged by the reaction time, the original sensitivity is restored in the light system following exposure to light in about one-third the time required for equilibrium to be reattained in the growth system.     

Effect of stimulus intensity on unit responses in the cat frontal cortex

Unit activity of the frontal cortex during changes in stimulus intensity in the near-threshold range (15–16 dB above the threshold for the combined evoked potential) was investigated by an extracellular recording method in acute experiments on cats anesthetized with chloralose (70 mg/kg). Comparative analysis of unit responses in specific (SI) and nonspecific projection areas revealed basically similar changes in pattern during an increase in stimulus intensity: A decrease in the latent period, an increase in the total frequency and the phasic character of the discharge, and an increase in the probability of response. However, a relatively stable latent period and probability of response were observed in specific projection neurons for a stimulus intensity of 3–5 threshold units, whereas for the nonspecific projection neurons it was observed for a stimulus intensity of 10–15 threshold units. All sensory projections in the frontal cortex are formed by two inputs: short-latency low-threshold and long-latency high-threshold. Analysis of modality-dependent differences in the threshold of sensitivity and the latent period of response of the polysensory neurons suggests that stimuli of different modalities converge directly on cortical neurons.A. A. Zhdanov Leningrad State University. Translated from Neirofiziologiya, Vol. 8, No. 6, pp. 606–612, November–December, 1976.