REACTION TO VISUAL FLICKER IN THE NEWT TRITURUS

The flicker response curve for the newt Triturus viridescens (water phase) has much the same quantitative structure as that found with various fresh-water teleosts at the same temperature (21.5°). The variability of critical intensity and of critical flash frequency likewise follows the same rules. The cone portion of the F - log I curve is much more widely spread, however. This, and the rather low maximum to which the rod curve rises, produce a considerable overlapping of the two parts additively fused. In addition, and to an extent which differs in various individuals, there is apparent a slight departure from the probability integral form of the cone curve. Reasons are given for considering that this is possibly connected with the role of an additional (small) number of (perhaps temporary, or developmental) retinal elements in addition to the typical rods and cones.     

TEMPERATURE AND CRITICAL ILLUMINATION FOR REACTION TO FLICKERING LIGHT : IV. ANAX NYMPHS

At fixed flash frequency (_F = 20, _F = 55) and with constant light time fraction (50 per cent) in the flash cycle, the critical illumination I for response of Anax nymphs to visual flicker falls continuously as the temperature rises. The temperature characteristic µ for the measure of excitability (1/I) increases continuously with elevation of temperature. The form of the F - log I curve does not change except at quite high temperature (35.8°), and then only slightly (near F = 55); F_max. is not altered. The very unusual form of the 1/I curve as a function of temperature is quantitatively accounted for if two processes, with respectively µ = 19,200 and µ = 3,400, contribute independently and simultaneously to the control of the speed of the reaction governing the excitability; the velocities of these two processes are equal at 15.9°.     

THE FLICKER RESPONSE CONTOUR FOR THE FROG

The flicker response contour for the frog Rana pipiens exhibits the duplex character typical for most vertebrates. By comparison (under the same conditions of temperature, 21.5°, and light-time fraction, = 0.5), the low intensity section of the F - log I curve is the smallest thus far found. The cone portion of the curve is satisfactorily described by a probability integral. The rod part represents the addition of a small group of sensory effects upon the lower end of the cone curve, from which it can be analytically separated. The relation between the two groups of sensory effects permits certain tests of the rule according to which (in homogeneous data) I_m and σ_1I1 are in direct proportion.     

TEMPERATURE CHARACTERISTICS FOR THE PRODUCTION OF CO2 BY GERMINATING SEEDS OF LUPINUS ALBUS AND ZEA MAYS

The rates of production of CO₂ by germinating seeds of Lupinus albus and Zea mays were studied between temperatures 12.5° and 25°C. with the HCl-Ba(OH)₂ titration method. The temperature characteristics found are different from those previously obtained for the oxygen consumption of the same seeds germinated in the same manner. For Lupinus, the temperature characteristics above and below the critical temperature of 20° are 16,100 ± and 24,000 ± calories respectively. For Zea, no evidence of a critical temperature was found in this region, and the temperature characteristic is 20,750 ± calories throughout the range of temperature tested. The possible interpretations of the difference in the values of temperature characteristics for oxygen consumption and for production of CO₂ are noted.     

TEMPERATURE AND CRITICAL ILLUMINATION FOR REACTION TO FLICKERING LIGHT : II. SUNFISH

The curve connecting mean critical illumination (I_m) and flicker frequency (F) for response of the sunfish Lepomis (Enneacanthus gloriosus) to flicker is systematically displaced toward lower intensities by raising the temperature. The rod and cone portions of the curve are affected in a similar way, so that (until maximum F is approached) the shift is a nearly constant fraction of I_m for a given change of temperature. These relationships are precisely similar to those found in the larvae of the dragonfly Anax. The modifications of the variability functions are also completely analogous. The effects found are consistent with the view that response to flicker is basically a matter of discrimination between effect of flashes of light and their after effects,—a form of intensity discrimination. They are not consistent with the stationary state formulation of the shape of the flicker curve. An examination of the relationships between the cone portion and the rod portion of the curves for the sunfish suggests a basis for their separation, and provides an explanation for certain "anomalous" features of human flicker curves. It is pointed out how tests of this matter will be made.     

THE FLICKER RESPONSE CURVE FOR FUNDULUS

After Fundulus heteroclitus have been for some time in the laboratory, under conditions favorable for growth, and after habituation of the fishes to the simple routine manipulations of the observational procedure required, they are found to give reproducible values of the mean critical flash illumination (I_m) resulting in response to visual flicker. The measurements were made with equality of light time and dark time in the flash cycle, at 21.5°C. Log I_m as a function of flash frequency F has the same general form as that obtained with other fishes tested, and for vertebrates typically: the curve is a drawn-out S, with a second inflection at the low I end. In details, however, the curve is somewhat extreme. Its composite form is readily resolved into the two usual parts. Each of these expresses a contribution in which log I, as a function of F, is accurately expressed by taking F as the summation (integral) of a probability distribution of d log I, as for the flicker response contour of other animals. As critical intensity I increases, the contribution of rod elements gradually fades out; this decay also adheres to a probability integral. The rod contribution seen in the curve for Fundulus is larger, absolutely and relatively to that from the cones, than that found with a number of other vertebrates. The additive overlapping of the rod and cone effects therefore produces a comparatively extreme distortion of the resulting F-log I curve. The F-log I_m curve is shifted to lower intensities as result of previous exposure to supranormal temperatures. This effect is only very slowly reversible. The value of F _max. for each of the components of the duplex curve remains unaffected. The rod and cone segments are shifted to the same extent. The persisting increase of excitability thus fails to reveal any chemical or other differentiation of the excitability mechanism in the two groups of elements. Certain bearings of the data upon the theory of the flicker response contour are discussed, with reference to the measurements of variation of critical intensity and to the form of the F-log I curve. The quantitative properties of the data accord with the theory derived from earlier observations on other forms.     

CRITICAL ILLUMINATION AND FLICKER FREQUENCY IN RELATED FISHES

Flicker response curves have been obtained at 21.5°C. for three genera of fresh water teleosts: Enneacanthus (sunfish), Xiphophorus (swordtail), Platypoecilius (Platy), by the determination of mean critical intensities for response at fixed flicker frequencies, and for a certain homogeneous group of backcross hybrids of swordtail x Platy (Black Helleri). The curves exhibit marked differences in form and proportions. The same type of analysis is applicable to each, however. A low intensity rod-governed section has added to it a more extensive cone portion. Each part is accurately described by the equation F = F_max./(1 + e ^{-p log-p logI/I_i}), where F = flicker frequency, I = associated mean critical intensity, and I_i is the intensity at the inflection point of the sigmoid curve relating F to log I. There is no correlation between quantitative features of the rod and cone portions. Threshold intensities, p, I_i, and F_max. are separately and independently determined. The hybrid Black Helleri show quantitative agreement with the Xiphophorus parental stock in the values of p for rods and cones, and in the cone F_max.; the rod F_max. is very similar to that for the Platy stock; the general level of effective intensities is rather like that of the Platy form. This provides, among other things, a new kind of support for the duplicity doctrine. Various races of Platypoecilius maculatus, and P. variatus, give closely agreeing values of I_m at different flicker frequencies; and two species of sunfish also agree. The effect of cross-breeding is thus not a superficial thing. It indicates the possibility of further genetic investigation. The variability of the critical intensity for response to flicker follows the rules previously found to hold for other forms. The variation is the expression of a property of the tested organism. It is shown that, on the assumption of a frequency distribution of receptor element thresholds as a function of log I, with fluctuation in the excitabilities of the marginally excited elements, it is to be expected that the dispersion of critical flicker frequencies in repeated measurements will pass through a maximum as log I is increased, whereas the dispersion of critical intensities will be proportional to I_m; and that the proportionality factor in the case of different organisms bears no relation to the form or position of the respective curves relating mean critical intensity to flicker frequency. These deductions agree with the experimental findings.     

ON THE TEMPERATURE CHARACTERISTICS FOR FREQUENCY OF BREATHING MOVEMENTS IN INBRED STRAINS OF MICE AND IN THEIR HYBRID OFFSPRING. I

Young mice of a selected line of the dilute brown strain of mice exhibit over the range 15–25°C. (body temperature) a relation of frequency of breathing movements to temperature such that when fitted by the Arrhenius equation the data give a value for the constant µ of 24,000± calories or, less frequently, 28,000±. Young mice of an inbred albino strain show over the range 15–20°C. a value of µ = 34,000±, or, less frequently, 14,000±, with a critical temperature at about 20°C. and a value of µ = 14,000± above 20°C. The F₁ hybrids of these two strains, and the backcross generations to either parent strain, exhibit only those four values of the temperature characteristic observed in the parent strains and none other. One may therefore speak of the inheritance of the value of the constant µ, but the inheritance shows in this instance no Mendelian behavior. Furthermore there appears to be inherited the occurrence (or absence) of a critical temperature at 20°C. These experiments indicate the "biological reality" of the temperature characteristics.     

TEMPERATURE CHARACTERISTICS FOR METABOLISM OF CHLORELLA : I. THE RATE OF O2 UTILIZATION OF CHLORELLA PYRENOIDOSA WITH ADDED DEXTROSE

The temperature characteristic for the rate of O₂ consumption by Chlorella pyrenoidosa suspended in Knop solution containing 1 per cent glucose was studied between 1° and 27°C. with the Warburg technic. The value of µ was found to be about 19,000 ±1,000 cal. There is some indication of a critical temperature at 20°C., with shift to a lower µ above this temperature. The effect of sudden changes in temperature on the rate of respiration and the variation of the latter with time at constant temperatures are discussed. It is concluded that the "normal" respiration (in absence of external glucose) does not appear in the determination of this temperature characteristic.     

TEMPERATURE CHARACTERISTICS FOR PULSATION FREQUENCY IN GONIONEMUS

The frequency of contraction of the bell of Gonionemus was studied in relation to temperature, with intact animals and also where different operations were made on the nervous system. A number of values of µ are found for intact animals namely 8,100±, 10,500±, 32,000± and 22,500±, with critical temperatures at 9.6°, 12.3°, and 14.0°. Four different classes of operations were used: (1) Animals where the nerve ring was cut on two opposite sides of the bell; the µ values found are 10,500± and 21,300±, with a critical temperature at 13.4°. (2) Animals with four cuts through the nerve ring gave µ = 10,600 ± and µ = 21,000, with a critical temperature at 13.1°. (3) In animals where the bell was cut in half the temperature characteristic was found to be 16,900. And finally (4) in the animals where the nerve ring was totally removed µ values of 8,100, 16,000±, and 29,000 were found, with critical temperatures at 15.0° and 9.4°. These results are discussed from the standpoint of the theory which supposes that definite "temperature characteristics" may be associated with the functional activity of particular elements in a complex functional unit, and that these elements may be separately studied and identified by suitable experimental procedures involving the magnitudes of the respective temperature characteristics and the locations of associated critical temperatures. The swimming bell of medusæ with its marginal sense organs permits a fairly direct approach to such questions. It is found that even slight injuries to the marginal nerve ring, for example, produce specific modifications in the temperature relations which are different from those appearing when the organism is cut in half.     

CRITICAL ILLUMINATION AND FLICKER FREQUENCY,AS A FUNCTION OF FLASH DURATION: FOR THE SUNFISH

From the relations between critical illumination in a flash (I_m) and the flash frequency (F) for response of the sunfish to visual flicker when the proportion of light time to dark time (t_L/t_D) in a flicker cycle is varied at one temperature (21.5°) the following results are obtained: At values of t_L/t_D between 1/9 and 9/1 the F - log I_m curves are progressively shifted toward higher intensities and lower F_max.. F_max. is a declining rectilinear function of the percentage of the flash cycle time occupied by light. The rod and the cone portions of the flicker curve are not shifted to the same extent. The cone portion and the rod region of the curve are each well described by a probability integral. In terms of F as 100 F/F_max. the standard deviation of the underlying frequency distribution of elemental contributions, summed to produce the effect proportional to F, is independent of t_L/t_D. The magnitude of log I_m at the inflection point (r''), however, increases rectilinearly with the percentage light time in the cycle. The proportionality between I_m and σ_II1 is independent of t_L/t_D. These effects are interpreted as consequences of the fact that the number of elements of excitation available for discrimination of flicker is increased by increasing the dark interval in a flash cycle. Decreasing the dark interval has therefore the same kind of effect as reducing the visual area, and not that produced by decreasing the temperature.     

TEMPERATURE CHARACTERISTICS FOR THE OXYGEN CONSUMPTION OF GERMINATING SEEDS OF LUPINUS ALBUS AND ZEA MAYS

The rate of oxygen consumption by germinating seeds of Lupinus albus and of Zea mays was studied as a function of temperature (7–26°C.). The Warburg manometer technique was used, with slight modifications. Above and below a critical temperature at 19.5°C. the temperature characteristic for oxygen consumption by Lupinus albus was found to be µ = 11,700± and 16,600 respectively. The same critical temperature was encountered in the case of Zea mays, with temperature characteristics µ = 13,100± above and µ = 21,050 below that temperature.     

TEMPERATURE AND CRITICAL ILLUMINATION FOR REACTION TO FLICKERING LIGHT : V. XIPHOPHORUS,PLATYPOECILIUS, AND THEIR HYBRIDS

For the teleosts Xiphophorus montezuma, Platypoecilius maculatus, and their F ₁ hybrids the temperature characteristics (µ in Arrhenius'' equation) are the same for the shift of the low intensity and the high intensity segments of the respective and different flicker response contours (critical intensity I as a function of flash frequency F, with light time fraction constant, at 50 per cent). The value of µ is 12,500 calories or a very little less, over the range 12.5 to 36°. This shows that 1/I can be understood as a measure of excitability, with F fixed, and that the excitability is governed by the velocity of a chemical process common to both the classes of elements represented in the duplex performance curve (rods and cones). It is accordingly illegitimate to assume that the different shapes of the rod and cone branches of the curves are determined by differences in the chemical mechanisms of excitability. It is also forbidden to assume that the differing form constants for the homologous segments in the curves for two forms (X. and P.) are the reflections of a difference in the chemical factors of primary excitability. These differences are determined by statistical factors of the distribution of excitabilities among the elements implicated in the sensory effect vs. intensity function, and are independent of temperature and of the temperature characteristic. It must be concluded that the physicochemical nature of the excitatory process cannot be deduced from the shape of the performance contour. The form constants (σ''_{log I} and F_max.) for F vs. log I are specifically heritable in F ₁, although µ is here the same as for X. and P. In an intergeneric cross one cannot in general expect Mendelian simplicity of segregation in subsequent generations, and in the present case we find that F ₂ individuals are indistinguishable from F ₁, both as regards F vs. log I and as regards the variation of I within a group of 17 individuals. The result in F ₂ definitely shows, however, that certain specific statistical form constants for the F-log I contour are transmissible in inheritance. It is pointed out that there thus is provided an instance in which statistical (distribution) factors in performance characteristics involving the summating properties of assemblages of cellular units are heritable in a simple manner without the implication of detectable differences in chemical organization of the units involved. This has an important bearing upon the logic of the theory of the gene.     

INTERMITTENT STIMULATION BY LIGHT : III. THE RELATION BETWEEN INTENSITY AND CRITICAL FUSION FREQUENCY FOR DIFFERENT RETINAL LOCATIONS

When measurements of the critical fusion frequency for white light over a large range of intensities are made with the rod-free area of the fovea, the relation between critical frequency and log I is given by a single sigmoid curve, the middle portion of which approximates a straight line whose slope is 11.0. This single relation must be a function of the foveal cones. When the measurements are made with a retinal area placed 5° from the fovea, and therefore containing both rods and cones, the relation between critical frequency and log I shows two clearly separated sections. At the lower intensities the relation is sigmoid and reaches an upper level at about 10 cycles per second, which is maintained for 1.25 log units, and is followed by another sigmoid relationship at the higher intensities similar to the one given by the rod-free area alone. These two parts of the data are obviously separate functions of the rods at low intensities and of the cones at high intensities. This is further borne out by similar measurements made with retinal areas 15° and 20° from the fovea where the ratio of rods to cones is anatomically greater than at 5°. The two sections of the data come out farther apart on the intensity scale, the rod portion being at lower intensities and the cone portion at higher intensities than at 5°. The general form of the relation between critical frequency and intensity is therefore determined by the relative predominance of the cones and the rods in the retinal area used for the measurements.     

STIMULATION OF FUNDULUS BY OXALIC AND MALONIC ACIDS AND BREATHING RHYTHM AS FUNCTIONS OF TEMPERATURE

1. Chemical stimulation as a function of temperature was studied by using oxalic acid in fresh and salt water and malonic acid in salt water as stimulating agents on Fundulus. According to the Arrhenius equation the following µ values were obtained for the various acid solutions between 0 and 29°C.: for 0.002N oxalic in fresh water—15,800; 33,000; for 0.0008N oxalic in fresh water—15,800; 33,000; 48,000; for 0.002N oxalic in salt water—19,400; 24,100; 56,500; for 0.004N and 0.002N malonic in salt water—20,600; 65,000. At a critical temperature there is a sharp transition from one thermal increment to another. 2. The chemical processes controlling stimulation do not change with concentration, for different normalities of a single acid yield the same µ values. Distinctly different temperature characteristics were obtained for stimulation by oxalic in salt and fresh water. Likewise stimulation by oxalic and malonic in salt water yielded very different increments. This temperature study indicates that the controlling chemical reactions determining rate of response are different for the same acid in two different environments, or for two dibasic acids in the same environment. Other work indicates, however, that the fundamental stimulation system is the same for all the adds in both environments. Chemical rather than physical processes limit the rate of response since all the values are above 15,000. Stimulation depends upon a series of interrelated chemical reactions, each with its own temperature characteristic. Under varying conditions (e.g. change of temperature, environment, or acid) different chemical reactions may become the slowest or controlling process which determines the rate of response. 3. The variation of response, as measured by the probable error of the mean response time of the fish, is the same function of temperature as reaction time itself. Hence variability is not independent of reaction time and is controlled by the same catenary series of events which determine rate of response to stimulation. 4. Breathing rhythm of Fundulus as related to temperature was studied in both salt and fresh water. In salt water the temperature characteristic is 8,400 while in fresh water it is 16,400 below 9.5°C., and 11,300 above this critical temperature. These µ values are typical of those which have been reported by other workers for respiratory and oxidative biological phenomena. A change in thermal increment with an alteration in environment indicates that different chemical reactions with characteristic velocity constants are controlling the breathing rhythm in salt and fresh water.     

TEMPERATURE CHARACTERISTIC FOR PRODUCTION OF CO2 BY PHASEOLUS SEEDLINGS

The temperature characteristic for respiratory production of CO₂ by young seedlings of Phaseolus aureus (Roxb.) is µ = 16,500 calories, 12–21°C., even when the analyses depend upon the use of many seedlings crowded in a small respiration chamber, provided reasonable precautions are taken to avoid injury and to permit proper thermal adaptation. There is evidence of a definite critical temperature at 20–21°. These findings agree quantitatively with those obtained with other similar seedlings, and contradict the results reported by Kurbatov and Leonov (1930); the reasons for this are analysed.     

Diapause Induction and Termination in the Small Brown Planthopper,Laodelphax striatellus (Hemiptera: Delphacidae)

The small brown planthopper, Laodelphax striatellus (Fallén) enters the photoperiodic induction of diapause as 3rd or 4th instar nymphs. The photoperiodic response curves in this planthopper showed a typical long-day response type with a critical daylength of approximately 11 h at 25°C, 12 h at 22 and 20°C and 12.5 h at 18°C, and diapause induction was almost abrogated at 28°C. The third stage was the most sensitive stage to photoperiod. The photoperiodic response curve at 20°C showed a gradual decline in diapause incidence in ultra-long nights, and continuous darkness resulted in 100% development. The required number of days for a 50% response was distinctly different between the short- and long-night cycles, showing that the effect of one short night was equivalent to the effect of three long nights at 18°C. The rearing day length of 12 h evoked a weaker intensity of diapause than did 10 and 11 h. The duration of diapause was significantly longer under the short daylength of 11 h than it was under the long daylength of 15 h. The optimal temperature for diapause termination was 26 and 28°C. Chilling at 5°C for different times did not shorten the duration of diapause but significantly lengthened it when chilling period was included. In autumn, 50% of the nymphs that hatched from late September to mid-October entered diapause in response to temperatures below 20°C. The critical daylength in the field was between 12 h 10 min and 12 h 32 min (including twilight), which was nearly identical to the critical daylength of 12.5 h at 18°C. In spring, overwintering nymphs began to emerge in early March-late March when the mean daily temperature rose to 10°C or higher.     

THE TEMPERATURE CHARACTERISTIC OF RESPIRATION OF AZOTOBACTER

The temperature characteristic of respiration of Azotobacter vinelandii possesses a constant value of 19,330 ± 165 over the temperature range 20–30°C. This value is independent of pH, oxygen tension, age of culture, and other factors within the limits studied. The optimum temperature of respiration is 34–35°C., with limits at about 10° and 50°C.