Frequency as a releaser in the courtship song of two crickets,Gryllus bimaculatus (de Geer) and Teleogryllus oceanicus: a neuroethological analysis

1. The courtship behavior of male field crickets, Gryllus bimaculatus (De Geer) and Teleogryllus oceanicus, is a complex, multimodal behavioral act that involves acoustic signals (a courtship song; Fig. 1A,B). The dominant frequency is 4.5 kHz for T. oceanicus song (Fig. 1A) and 13.5 kHz for G. bimaculatus (Fig. IB).
2. When courting males are deprived of their courtship song by wing amputation, their courtship success declines markedly but is restored when courting is accompanied by tape-recordings of their courtship songs or a synthetic courtship song with only the dominant frequency of the natural song; other naturally occurring frequency components are ineffective for restoring mating success (Figs. 4, 5).
3. It has been suggested that an identified auditory interneuron, AN2, plays a critical role in courtship success. Chronic recordings of AN2 in an intact, tethered female show that AN2's response to the natural courtship song and synthesized songs at 4.5 and 13.5 kHz is similar in T. oceanicus. By contrast, in G. bimaculatus, AN2's response to the natural courtship song and synthesized song at 13.5 kHz, but not at 4.5 kHz, is similar (Figs. 2,3).
4. In behavioral experiments, playback of a 30 kHz synthetic courtship song in G. bimaculatus does not restore courtship success, yet this same stimulus elicits as strong a response from AN2 as does the normal courtship song (Fig. 6). Thus, contrary to earlier work by others, we conclude AN2 is not, by itself, a critical neural link in the courtship behavior of these two species of crickets.

     

Morphological and physiological properties of pheromone-triggered flipflopping descending interneurons of the male silkworm moth,Bombyx mori

1. The morphology of descending interneurons (DNs) which have arborizations in the lateral accessory lobe (LAL) of the protocerebrum, the higher order olfactory center, and have an axon in the ventral nerve cord (VNC), were characterized in the male silkworm moth, Bombyx mori.
2. Two clusters (group I, group II) of DNs which have arborizations mainly in the LALs were morphologically characterized. The axons of these DNs are restricted to the dorsal part of the each connective (Figs. 1–5).
3. Pheromonal responses of the group I and group II DNs were characterized. Flipflopping activity patterns, which have two distinct firing frequencies (high and low) in response to sequential pheromonal stimulation, were usually recorded (Figs.6–10).
4. Two types of flipflopping activity patterns were classified into those that had an antiphasic relationship (called the ‘FF’ type) between the left and right connectives and those with a synchronized relationship (‘ff’ type) (Figs. 8–12). We propose that some group II DNs show ‘FF’ flipflopping activity patterns (Fig. 10).
5. A state transition was usually elicited by less than 10 ng bombykol, the principal pheromone component. Extra impulses were elicited during constant light stimulation (Fig. 9).
6. Our results suggest that the LAL olfactory pathways might be important for producing flipflopping activity patterns (Fig. 11).

     

Response characteristics of wide-field non-habituating non-directional motion detecting neurons in the optic lobe of the locust,Locusta migratoria

1. Extracellular recordings from wide-field nonhabituating non-directional (ND) motion detecting neurons in the second optic chiasma of the locust Locusta migratoria are presented. The responses to various types of stepwise moving spot and bar stimuli were monitored (Fig. 1)
2. Stepwise motion in all directions elicited bursts of spikes. The response is inhibited at stimulus velocities above 5°/s. At velocities above 10°/s the ND neurons are slightly more sensitive to motion in the horizontal direction than to motion in the vertical direction (Fig. 2). The ND cells have a preference for small moving stimuli (Fig. 3).
3. The motion response has two peaks. The latency of the second peak depends on stimulus size and stimulus velocity. Increasing the height from 0.1 to 23.5° of a 5°/s moving bar results in a lowering of this latency time from 176 to 130 ms (Fig. 4). When the velocity from a single 0.1° spot is increased from 1 to 16°/s, the latency decreases from 282 to 180 ms (Figs. 5–6).
4. A change-of-direction sensitivity is displayed. Stepwise motion in one particular direction produces a continuous burst of spike discharges. Reversal or change in direction leads to an inhibition of the response (Fig. 7).
5. It shows that non-directional motion perception of the wide-field ND cells can simply be explained by combining self-and lateral inhibition.

     

Wind-evoked evasive responses in flying cockroaches

1. A standing cockroach (Periplaneta americana) responds to the air displacement made by an approaching predator, by turning away and running. The wind receptors on the cerci, two posterior sensory appendages, excite a group of ventral giant interneurons that mediate this response. While flying, these interneurons remain silent, owing to strong inhibition; however, the dorsal giant interneurons respond strongly to wind. Using behavioral and electromyographic analysis, we sought to determine whether flying cockroaches also turn away from air displacement like that produced by an approaching flying predator; and if so, whether the cerci and dorsal giant interneurons mediate this response.
2. When presented with a wind puff from the side, a flying cockroach carries out a variety of maneuvers that would cause a rapid turn away and perhaps a dive. These are not evoked if the cerci are ablated (Figs. 4, 5, 6).
3. This evasive response appears to be mediated by a circuit separate from that mediating escape when the cockroach is standing (Fig. 7).
4. The dorsal giant interneurons respond during flight in a directional manner that is suited to mediate this behavior (Fig. 8).
5. Recordings of the wind produced by a moving model predator (Fig. 9), together with measurements of the behavioral latency of tethered cockroaches, suggest that the evasive response would begin just milliseconds before a predator actually arrives. However, as explained in the Discussion section, under natural conditions, the evasive response may well begin earlier, and could indeed be useful in escaping from predators.
6. If cockroaches had a wind-mediated yaw-correcting behavior, as locusts have, this could conflict with the wind-evoked escape. In fact, cockroaches show the opposite, yaw-enhancing response, mediated by the cerci, that does not present a conflict with escape (Figs. 10–14).

     

Multisensory control of escape in the cockroach Periplaneta americana

1. All giant interneurons (GIs) were ablated from the nerve cord of cockroaches by electrocautery, and escape behavior was analyzed with high-speed videography. Animals with ablations retained the ability to produce wind-triggered escape, although response latency was increased (Table 1, Fig. 4). Subsequent lesions suggested that these non-GI responses depended in part on receptors associated with the antennae.
2. Antennal and cereal systems were compared by analyzing escape responses after amputating either cerci or antennae. With standard wind stimuli (high peak velocity) animals responded after either lesion. With lower intensity winds, animals lost their ability to respond after cereal removal (Fig. 6).
3. Removal of antennae did not cause significant changes in behavioral latency, but in the absence of cerci, animals responded at longer latencies than normal (Fig. 7).
4. The cercal-to-GI system can mediate short latency responses to high or low intensity winds, while the antennal system is responsive to high intensity winds only and operates at relatively longer latencies. These conclusions drawn from lesioned animals were confirmed in intact animals with restricted wind targeting the cerci or antennae only (Fig. 9).
5. The antennae do not represent a primary wind-sensory system, but may have a direct mechanosensory role in escape.

     

Time course of the Houseflies' landing response

The landing response of stationary flying houseflies Musca domestica has been recorded on video tape. The leg movements were quantitatively evaluated. It could be demonstrated that:

1. only the first two pairs of legs are involved in the reaction (Fig. 1). Prothoracic tarsi are lifted beyond the head, mesothoracic tarsi are lowered and moved sidewards (Fig. 2a and b).
2. the movement of the tarsal tips is mainly due to an opening of one single joint per leg, i.e. the femurtibia joint of the prothoracic leg (Fig. 2c), and the coxa-femur joint of the mesothoracic leg.
3. the landing reaction is a fixed action pattern which does not seem to require further sensory input once it is released (Fig. 4d).
4. the landing responses to a light-off stimulus and to expanding patterns with different angular velocities are indistinguishable (compare Fig. 3a-c with Fig. 2a-c). The only parameter that is obviously dependent on the stimulus conditions, is the latency of the reaction (Fig. 4a-c).

     

Polarization sensitivity in crayfish lamina monopolar neurons

1. Polarization sensitivity (PS) was examined in photoreceptors and lamina monopolar cells (LMCs) in two species of crayfish, Procambarus clarkii and Pacifasticus leniusculus. The measurements were made with intracellular recordings and broad field illumination.
2. PS is about 40% greater in Pacifasticus than in Procambarus (Table 1). In both species the LMC stationary PS profiles (estimated with flashes) are similar to those of receptors (Figs. 1 and 2). Both receptor and LMC sensitivity profiles are well described by cos² θ functions (Fig. 3). PS was observed in all receptors and 78% of LMCs.
3. When stimulated with a rotating polarizer, receptors and LMCs exhibit membrane potential modulation with phase predicted by the stationary PS profile (Fig. 5). In photoreceptors, the polarization-elicited percent modulation falls off steeply as intensity increases. The LMC modulation is stronger than that in receptors and relatively insensitive to the mean intensity (Figs. 6 to 8). For low intensities the LMC modulation is 100%. The LMC dynamic behavior is consistent with either an opponency mechanism or strong but polarization-insensitive lateral inhibition.
4. Receptors and LMCs exhibit steady-state differential sensitivity to stationary e-vector orientation (Fig. 9).
5. About 10% of the LMC neurons exhibit PS maxima separated by 90°. These results imply a nonlinear summation of signals from orthogonal receptor channels (Fig. 10).

     

Selective phonotaxis to advertisement calls in the gray treefrog Hyla versicolor: behavioral experiments and neurophysiological correlates

1. The significance of particular acoustic properties of advertisement calls for selective phonotaxis by the gray treefrog, Hyla versicolor (= HV), was studied behaviorally and neurophysiologically. Most stimuli were played back at 85 dB SPL, a level typically measured at 1–2 m from a calling male.
2. Females preferred stimuli with conspecific pulse shapes at 20° and 24°C, but not at 16°C. Tests with normal and time-reversed pulses indicated the preferences were not influenced by the minor differences in the long-term spectra of pulses of different shape.
3. Pulse shape and rate had synergistic or antagonistic effects on female preferences depending on whether the values of one or both of these properties in alternative stimuli were typical of those in HV or heterospecific (H. chrysoscelis = HC) calls.
4. More auditory neurons in the torus semicircularis were temporally selective to synthetic calls (90%) than to sinusoidally AM tones and noise (< 70%).
5. Band-pass neurons were tuned to AM rates of 15–60 Hz. Neurons were more likely to be tuned to HV AM rates ( < 40 Hz) when stimuli had pulses with HV rather than HC shapes.
6. Sharp temporal tuning was uncommon and found only in neurons with band-pass or low-pass characteristics.
7. Many neurons differed significantly in response to HV and HC stimulus sets. Maximum spike rate was more often elicited by an HV stimulus (74%) than by an HC stimulus (24%).
8. Differences in spike rates elicited by HV and HC stimuli were attributable to combinations of differences in the rise times and shapes of the pulses.

     

Physiological and pharmacological properties of responses to GABA and ACh by abdominal motor neurons in Manduca sexta

1. GABA, ACh, and other agents were applied by pressure ejection to the neuropil of the third abdominal ganglion in the isolated nerve cord of Manduca sexta. Intersegmental muscle motor neurons with dendritic arborizations in the same hemiganglion were inhibited by GABA (Fig. 2) and excited by ACh (Fig. 5).
2. Picrotoxin was a potent antagonist of GABA (Fig. 4A). Bicuculline reduced GABA responses in some motor neurons (Fig. 4C), but had no effect on many other motor neurons. Curare reduced ACh responses (Fig. 6A). Bicuculline was an effective ACh antagonist in most motor neurons tested (Fig. 6B).
3. Motor neurons with dendrites across the ganglion from the ejection pipette exhibited different responses to GABA and ACh. Contralateral motor neurons often showed smaller, delayed hyperpolarizing GABA responses (Fig. 7). On two occasions, contralateral motor neurons had excitatory responses (Fig. 8). Contralateral motor neurons were hyperpolarized by ACh (Fig. 9). The inhibitory responses had only slightly longer latencies than ipsilateral excitatory ACh responses (Fig. 10A). The contralateral inhibitory ACh responses, but not the ipsilateral excitatory ACh responses, were eliminated by TTX (Fig. 10B).
4. A model, which includes inhibitory interneurons that cross the ganglionic midline to inhibit their contralateral homologs and motor neurons (Fig. 11), is proposed to account for contralateral responses to GABA and ACh and antagonistic patterns of activity of motor neurons during mechanosensory reflex responses.

     

Oxygen consumption and respiratory behaviour of three Nile fishes

1. The respiratory behaviour and patterns of oxygen consumption of three Nile species have been investigated.
2. Tilapia nilotica showed a typical pattern of oxygen consumption with an ambient region, adaptive plateau and lethal region (Fig. 2).
3. Specimens of Polypterus senegalus and Clarias lazera (body weights 20–30 and 30–45 g respectively) showed patterns of consumption comparable to that of Tilapia (Fig. 3a and 4a). In larger specimens of the two species the adaptive plateau was either insignificant or completely absent.
4. Specimens of Polypterus and Clarias (20–30 g and 30–45 g respectively) could survive in waters saturated with oxygen (7.4 mg/l) but their tolerance to lower oxygen concentrations was limited. Larger specimens of Polypterus and Clarias failed to survive in oxygen saturated waters.
5. The tolerance of Tilapia nilotica to extremely low oxygen concentration is an adaptation of a tropical and completely aquatic species. Polypterus and Clarias resort to their compensatory mechanisms only when the aquatic respiratory surface fails to satisfy their oxygen requirements.

     

A study on the rate of water transport of the clam katelysia opima in relation to environmental conditions

1. The neutral red technique was employed to study the rate of filtration in Katelysia opima.
2. The weight specific water filtration was found to be greater for younger clams compared to the older ones.
3. The rate of water filtration increased with decreasing salinity.
4. Water filtration was found to increase as temperature increased, reaching a maximum at 35°C. but then sharply decreasing at 39°C.
5. Light had no significant effect on the rate of filtration.
6. Suspended matter was found to affect the rate of water filtration.
7. The rate of filtration was low at high pH and high in low pH.
8. The rate of water filtration was found to be faster during high tide than during low tide.
9. The presence of the parasitic crab, Pennotheris sp., in the mantle cavity of clams had a marked effect on the particle filtration.
10. Accidental cut of the siphon tips had no effect on the rate of filtration.

     

Effects of Environmental Perturbations on Short-Term Phytoplankton Production off Lawson's Bay,A Tropical Coastal Embayment

1. The phytoplankton cycle off Lawson's Bay, Waltair follows a bimodal pattern with a major peak during March–May; a minor peak during October–November months and with a low production during the summer months i.e., June–August.
2. During the summer months of 1957, 1958, 1960 and 1962 dumping of dredged spoil from the entrance channel of the harbour into the sea resulted in a natural enrichment of waters.
3. Following this enrichment, there was a qualitative and quantitative increase in the phytoplankters thus leading to the development of a bloom.
4. Only Thalassiosira subtilis and Chaetoceros curvisetus commonly bloomed during the four years.
5. The increase in gross production which varied from 3–13 fold and the high photosynthesis-respiration ratios 5.1 to 10.5 indicated that the bloom populations were in a healthy state.
6. The decrease of the populations to the initial levels suggests that some unknown factor, other than those investigated must have been operating.
7. Consequences of eutrophication of different origins on stimulation of phytoplankton production are briefly discussed.

     

Basic processes of locomotor coordination in the rock lobster

Rock lobsters are able to perform long and stereotyped stepping sequences above a motor driven treadmill. Forward walking samples are estimated by mean of statistical methods to draw out the basic rules involved in the locomotor behaviour (Fig. 1).

- The spatial and temporal parameters defined in a single propulsive leg are either invariable in respect to the imposed speed, as the mean step length (L), the return stroke duration (Tr) and the pause times (T's, T'r), or speed dependent as the power stroke duration (Ts) and the whole period (Figs. 2 and 3).

- The interleg phase coupling is strong and stable in the ipsilateral rear pairs (4–5), these legs acting most of the time in absolute coordination (1:1) or in harmonic ratio (2:1). In the contralateral pairs (R4-L4, R5-L5) the legs roughly operate in antiphase, but the relationship appears much weaker and variable, with frequent episodes of relative coordination (Fig. 4).

- The time intervals between the ground contact of any leg and the swing initiation in the nearest ones appear somewhat constant and could be closely related to the mechanism of stepping synchronization. The “5 on - 4 off” delay, very stable and always positive, suggests that the rear legs could exert a predominant influence upon the rhythmical movements of the next anterior ipsilateral appendages (Fig. 5).

- To test the contralateral relationships, the treadmill belts can be decoupled in order to impose different walking speeds on each side. Such a conflicting stimulus reveals that:

1. The relative hierarchy always observed between the ipsilateral legs can be artificially created between the two sides (Fig. 6).
2. The driving influence of a given leg is closely linked to the intensity of EMG's discharges in its power stroke muscles.
3. The contralateral appendages are able to walk in absolute coordination despite a large speed difference between the two sides (up to 4 cm/s). Under such a constraint, the walking legs alter its invariable parameters (L and Tr) to reach a common step period and steadily maintain the alternating pattern (Figs. 6 and 7).

     

Growth of a floating aquatic weed,Salvinia under standard conditions

1. Growth of the floating aquatic weed, Salvinia, in sterile culture was exponential for at least 2 weeks under standardized conditions.
2. Increase in light intensity or in CO₂ resulted in increases in growth rate, but did not extend the exponential period of growth.
3. This aquatic plant, like many others, discriminates against calcium relative to strontium.
4. In culture Salvinia exhibited luxury consumption of N and P.
5. Because of high C/N ratios, Salvinia may not be a favorable source of animal food, but might be useful in nutrient removal schemes.
6. In sterile culture, S. molesta produced fewer leaves than S. minima, but maintained a significant increase in leaf area and dry weight. This may be correlated with the ability of the first species to rapidly spread over tropical waterways.

     

Encoding of plant odours by receptor neurons in the pine weevil (Hylobius abietis) studied by linked gas chromatography-electrophysiology

Receptor neuron responses to plant volatiles, trapped by head-space procedures, were examined in the pine weevil Hylobius abietis, using gas chromatography linked with electrophysiological recordings from single neurons. Seventy-two receptor neurons were tested 173 times for various plant volatile mixtures, either via a polar or a non-polar column.

1. All responses appeared as increased firing rates which followed the concentration profiles of the GC-eluted compounds.
2. The neurons were classified separately for the two column types in 17 and 19 groups respectively, according to the compounds they responded to. It suggests that the plant odour information is encoded by a large, but limited number of receptor neuron types.
3. Most neurons responded to a limited number of compounds (1–5) and showed a marked best response to one of them, whereas additional responses to several other components which seems to be structurally similar, was recorded for some neurons. It suggests that the plant odour receptor neurons are rather narrowly than broadly tuned, and that each neuron is specialized for receiving information about one or a few related compounds.
4. Most neurons responded to monoterpenes, whereas the other neurons responded to compounds of other categories.
5. Both major and minor plant volatile components activated specifically receptor neurons.

     

The effects of captivity on the electric organ discharge and plasma hormone levels in Gnathonemus petersii (Mormyriformes)

1. Experiment 1 employed a repeated measures design to examine the effects of captivity on sex differences in the electric organ discharge (EOD) of Gnathonemus petersii, newly imported from Africa, and maintained individually or in groups.
2. On the day of import, males exhibited longer durations of phases 2 and 3 of the EOD and lower peak power spectral frequencies (PPSFs) than females.
3. After 14 days in captivity in the laboratory, the sex differences were eliminated. After 37 days of captivity, all sex differences were still abolished, or even reversed depending on housing conditions. Males exhibited the most dramatic changes in EODs and females appeared to have higher testosterone (T) levels than males.
4. Experiment 2 was designed to investigate the effects of captivity on both behavior and endocrine status in 58 newly imported males. In this independent group design, EOD data and blood were collected from subjects over 15 days.
5. Decreases in phase 3 of the EOD and increases in PPSFs progressed over the 15 day experimental period, becoming statistically significant by days 10 and 15, respectively. Regardless of housing conditions, both T and 11-keto T dramatically decreased to near non-detectable levels by Day 5 in the laboratory.
6. Captivity causes rapid and profound changes in the endocrine system which result in dramatic changes in steroid-sensitive EODs. These findings directly link captivity, hormones, and behavior, and show why feral animals brought into captivity usually do not exhibit sexual behavior.

     

The oxygen consumption of mayfly (Ephemeroptera) and stonefly (Plecoptera) larvae at different oxygen concentration

1. The aim of this investigation was to elucidate how four acquatic insect larvae, from different habitats and having different respiratory organs or types of respiratory regulation, react to a lowered oxygen concentration, and how their oxygen consumption is affected. The species investigated were the stoneflies Taeniopteryx nebulosa, Diura nanseni and Nemoura cincerea and the mayfly Cloëon dipterum.
2. The measurements were performed in a respiratory apparatus of open, flowing-water type. Its design is shown in Fig. 1. Water of known oxygen concentration was allowed to flow past the experimental larvae. The oxygen consumption of the larvae was calculated from the lowering of the oxygen concentration in which ensued.
3. The water used in the experiments was standardized, so that the electrode had the necessary stability (conductivity 470 micromhos/cm). The calcium ion was excluded in order to prevent the precipitation of CaCO₃ in the electrode capillary.
4. A large variation in the values of oxygen consumption was found as seen in Fig. 2–5. The reason for that is a corresponding variation in the motor activity of the experimental animals.
5. The physiological reasons for the general form of the curves A and C in Fig. 2–5 are discussed. The curves A and C represent oxygen consumption of the larvae at different degrees of stimulation, entailing different levels of motor activity. Curve A represents intentinally activated animals, curve C non-activated, motionless animals. The curves A and C are boundary curves corresponding to a sort of scope for activity of the animals. Over this scope area a series of curves of the same form could in principal be construed, representing different degrees of stimulation.
6. Within a certain oxygen concentration interval a motor activation was observed caused by a reduced oxygen concentration. The result of that activation can be seen in Fig. 2–5 as a zone with no or very few oxygen consumption values between curve C and D. The more easily activated the species is, the broader the zone will be. Cloën has the most narrow zone and was observed to be less activated than the other species.
7. Small larvae of Cloën (2–4 mm and 42–6 mm) and Nemoura (2–4 mm) showed clearly a greater ability to take up oxygen at low oxygen concentrations than full-grown larvae (see Fig. 8 and 9).
8. The critical point on the curve representing mean oxygen consumption as a function of oxygen concentrations was found to be at 2–5 mg O₂/1 for Taeniopteryx and Diura, at 2.2–2.5 mg O₂/1 for Cloëon, and at about 2–7 mg O₂/1 for Nemoura. The values refer to 8°. Cloëon is the species which is exposed to the greatest variations in oxygen concentration in its natural environment.
9. No influence on the oxygen consumption of starvation for 4 to 5 days was found. No difference between the oxygen consumption values obtained in the presence or in the absence of calcium ions could be observed during the experiments (Fig. 10, 11).
10. The basic picture obtained in this investigation is a set of oxygen consumption values scattered between a curve connecting highest values obtained and a curve of the standard metabolism together with a zone in which the larvae are activated by reduced oxygen concentrations. This picture is presumed to be general in aquatic animals with a well developed motor activity.

     

Survival,osmoregulatory ability,and respiration of idotea chelipes (Crustacea,Isopoda) from lake veere in different salinities and temperatures

1. An ecological and physiological study ofI. chelipes from Lake Veere, The Netherlands, was made.
2. Both osmoregulatory capacity and survival decrease with increasing temperature as well as with decreasing salinity.
3. Respiration experiments suggest that the need of energy by osmoregulatory activity may be supplied at the cost of other physiological processes, at any rate at temperatures of 10°C and higher.
4. It may be expected that, if temperatures higher than 15°C and salinities lower than 8‰ coincide, the population ofI. chelipes will be affected negatively.

     

The effect of temperature on the consumption,absorption and conversion of food in Ophiocephalus punctatus bloch

1. The overall feeding rate is maximum at 28°C and decreases at 20°C and 33°C for all sizes.
2. The smaller fish are cold-sensitive. At 28°C and at 33°C the time course rate shows the same decline, whereas at 20°C the response was anomalous.
3. Absorption efficiency is independent of size and temperature.
4. Rate of absorption and hence rate of conversion is primarily conditioned by rate of feeding.
5. At 20°C the conversion efficiency is directly related to size whereas at 28°C and 33°C it is inversely related to size.

     

Emmetropization and optical development of the eye of the barn owl (Tyto alba)

1. We have studied the development of the refractive state in young barn owls (Tyto alba pratincola). Strikingly, the eyes had severe refractive errors shortly after lid opening (which occurred around day 14 after hatching; average from 6 owls: 13.83 ± 1.47 days). Refractive errors vanished in the subsequent one or two weeks (Fig. 1, Fig. 2).
2. Refractive errors did not differ by more than 1 diopter (D) in both eyes of an individual (Fig. 2). Thus, non-visual control of eye growth was sufficient to produce non-random refractions. However, visual input was finally required to adjust the optical system to emmetropia.
3. Using in-vivo A-scan ultrasonography of ocular dimensions (Fig. 4A), photokeratometric measurements of corneal radius of curvature (Fig. 4B), and frozen sections of excised eyes (Fig. 3), we developed paraxial schematic eye models which described age-dependent changes in ocular parameters and were applicable through the ages from lid opening to fledging (Table 1). A schematic eye for the adult barn owl (European subspecies: Tyto alba alba) is also provided. Eye sizes in an adult owl of the American (Tyto alba pratincola) and the European subspecies (T. alba alba) were similar despite of different body weights (500 g and 350 g, respectively).
4. The schematic eyes were used to test which ocular parameters might have caused the recovery from refractive errors. However, none of the ocular dimensions measured underwent obvious changes in their growth curves as visual input became available. Apparently, coordinated growth of several ocular components produced emmetropia.
5. From the schematic eye model, the developmental changes in image brightness and image magnification were calculated (Fig. 5). In barn owl eyes, image size was not quite as extreme as in the tawny owl or the great horned owl. However, the image was larger and the f/number was lower than in diurnal birds of comparable weight (pigeon, chicken). The observation supports a conclusion that image size is maximised in owls to permit a higher degree of photoreceptor convergence for higher light sensitivity at dusk while spatial acuity remains comparable to diurnal birds with smaller eyes.