Wide-band spectral tuning of heat receptors in the pit organ of the copperhead snake (Crotalinae)

Receptors located in the facial pit organ of certain species of snake signal the presence of prey. Infrared radiation is an effective stimulus suggesting that these receptors may be low-threshold temperature receptors. We recorded from the nerve innervating the pit organ of snakes belonging to the family of Crotalinae while stimulating the receptive area with well-defined optical stimuli. The objective was to determine the sensitivity of these receptors to a wide range (0.400-10.6 micro m) of optical stimuli to determine if a temperature-sensitive or photosensitive protein initiated signal transduction. We found that receptors in the pit organ exhibited a unique broad response to a wide range of electromagnetic radiation ranging from the near UV to the infrared. The spectral tuning of these receptors parallels closely the absorption spectra of water and oxyhemoglobin, the predominant chromophore in tissue. Our results support the hypothesis that these are receptors activated by minute temperature changes induced by direct absorption of optical radiation in the thin pit organ membrane.     

Investigation of the transduction mechanism of infrared detection in Melanophila acuminata: photo-thermal-mechanical hypothesis

Differential phase optical low coherence reflectometry (OLCR) was used to detect sub-wavelength displacements in the infrared-sensitive thoracic pit organ of Melanophila acuminata (Coleoptera: Buprestidae) upon absorption of infrared radiation at 3.39 microm. The displacement had more complex morphology but similar amplitude ( approximately 100 nm at 1 W cm(-2)) when compared to the displacement measured from the exocuticle in an alternate region on the beetle's body. In addition, a simplified finite difference model was developed to predict the temperature distribution and resultant thermal expansion in the pit organ tissue. The experimental and model results were interpreted to help clarify the mechanism by which the sensilla in the pit organ convert infrared radiation to neural signals. The results of this paper are discussed in relation to the photo-thermal-mechanical transduction hypothesis. This is the first experimental examination of the transduction mechanism in Melanophila acuminata.     

Dynamic sweating response of man to infrared irradiation in various spectral regions

In an attempt to detect differences in the thermal effect of infrared irradiation of different wavelengths, transient sweating response to infrared irradiation in various spectral regions was examined. In Series 1, the ventral or dorsal surface of the nude subject was irradiated repetitively for a period of 4 min (2 min on, 2 min off) by each of three kinds of infrared heaters with main emissivity in near-infrared (NIR; 0.7–2.8 m), intermediate-infrared (MIR; 1.5–5.8 m), and far-infrared (FIR; 2.8–25 m) regions. The sweating response on a non-irradiated area tended to be the greatest with MIR, while the magnitude of the sweating response on the irradiated area showed no consistent differences among various wavelengths. The results infer that MIR stimulated cutaneous thomoreceptors most effectively, while its direct effect on local sweat gland activity was minimal. In Series 2, the effects of 9–12 min irradiations in more restricted ranges of wavelength were compared by the combination of the three kinds of heaters with filters (translucent to wavelength ranges of 1.3–2.7, 2.7–3.5, 3.6–8.0 m, respectively). The sweating response on a remote area was predominantly greater with the range of 2.7–3.5 m than with the other wavelength ranges, while the local effect on sweating was minimal with this range. The results of Series 2 reinforce those of Series 1, indicating that the degree of stimulation of cutaneous thermoreceptors and of direct thermal effect on sweat gland activity differ with spectral regions incident on the skin, thus affecting local and remote effects on the sweating response.     

Neurophysiological and behavioural evidence for an olfactory function for the dorsal organ and a gustatory one for the terminal organ in Drosophila melanogaster larvae

Multicellular electrophysiological responses from the dorsal organ on the cephalic lobes of third instar Drosophila melanogaster larvae (wild-type Canton S) stimulated with a cold-trapped banana volatile extract showed that this structure has an olfactory function in the fruit fly. Responses of the dorsal organ were also recorded to constituents of the banana volatile extract as they eluted from a gas chromatographic column (GC-coupled dorsal organ electrophysiology). The active chemostimulants were identified as 2-heptanone, isoamyl alcohol, hexyl acetate, hexanol and hexyl butyrate by gas chromatography-coupled mass spectrometry. Applying the same recording system to the terminal organ sensilla, no responses were obtained to either the banana volatile bouquet or its constituents. By contrast, high frequency multicellular responses were recorded in response to touching the terminal organ with the gustatory stimuli KCl and grapefruit juice; responses were absent on similar stimulation of the dorsal organ with either NaCl or KCl. This suggests a role for olfaction by the dorsal organ and for gustation by the terminal organ in Drosophila larvae.In a 7-mm high wind tunnel with a thin 1.2% agar floor, the Drosophila larvae showed odour-conditioned upwind responses in an air stream of 0.1 m/s bearing banana volatiles. Drosophila larvae responded best to the odour of cut bananas. A 1:1 mixture of the banana odour constituents 2-heptanone and hexanol (at either 50 or 100 &mgr;g source dose each) proved as attractive as the known larval attractants propionic acid and isoamyl acetate on their own at 100 &mgr;g, whereas hexanol and 2-heptanone on their own at a 100 &mgr;g source dose were less attractive. The stronger behavioural response to the banana volatile bouquet and to the binary mixture serves to underline the multireceptor nature of the dorsal organ response to food odour in Drosophila.     

The Python pit organ: imaging and immunocytochemical analysis of an extremely sensitive natural infrared detector

The Python infrared-sensitive pit organ is a natural infrared imager that combines high sensitivity, ambient temperature function, microscopic dimensions, and self-repair. We are investigating the spectral sensitivity and signal transduction process in snake infrared-sensitive neurons, neither of which is understood. For example, it is unknown whether infrared receptor neurons function on a thermal or a photic mechanism. We imaged pit organs in living Python molurus and Python regius using infrared-sensitive digital video cameras. Pit organs were significantly more absorptive and/or emissive than surrounding tissues in both 3-5 microns and 8-12 microns wavelength ranges. Pit organs exhibited greater absorption/emissivity in the 8-12 microns range than in the 3-5 microns range. To directly test the relationship between photoreceptors and pit organ infrared-sensitive neurons, we performed immunocytochemistry using antisera directed against retinal photoreceptor opsins. Retinal photoreceptors were labeled with antisera specific for retinal opsins, but these antisera failed to label terminals of infrared-sensitive neurons in the pit organ. Infrared-receptive neurons were also distinguished from retinal photoreceptors on the basis of their calcium-binding protein content. These results indicate that the pit organ absorbs infrared radiation in two major atmospheric transmission windows, one of which (8-12 microns) matches emission of targeted prey, and that infrared receptors are biochemically distinct from retinal photoreceptors. These results also provide the first identification of prospective biochemical components of infrared signal transduction in pit organ receptor neurons.     

Sensitivity threshold and response characteristics of infrared detection in the beetle Melanophila acuminata (Coleoptera: Buprestidae)

The minimum detection threshold of the infrared sensitive beetle, Melanophila acuminata, was measured with a helium-neon laser that emitted light at a wavelength of 3.39 microm. Extracellular recordings were taken both at the pit organ responsible for detection and at the interganglionic connectives in the thorax of the beetle. At the pit organ, generator and action potentials from single neurons were measured with a sharpened tungsten electrode. At the connectives that linked the fused second meso-/metathoracic and prothoracic ganglia, compound action potentials were measured with a tungsten hook electrode that encircled the connective. The latter recordings confirmed conveyance of infrared information through specific pathways to rostrally-situated sites in the nervous system of the beetle. The 50% probability irradiance threshold at which action potentials were elicited from the receptor and connectives occurred at 17.3 and 14.6 mW/cm(2), respectively. In addition to sensitivity threshold, several other characteristics of the response were quantified including dependence of generator potential latency, generator potential duration, spike frequency, and spike latency on irradiance, dependence of response strength (spike count) on exposure time, and flicker fusion frequency. The ability to detect infrared radiation is rare in nature, and these results provide valuable information necessary to understand this unique sensitivity.     

Thermal modeling of snake infrared reception: evidence for limited detection range

For more than 40 years, information has circulated with regard to the sensitivity of infrared pit organs in both boid and crotaline snakes (pythons and pit vipers, respectively). The most often quoted sensitivity is 0.003 degrees C and this value is based on the work of Bullock and co-workers (1956). Missing from previous work was a quantitative model of radiation transfer that would report sensitivity not in terms of degrees Celsius, but rather sensing distance. Since prey detection is often cited as the function of the infrared pit organ, quantification of this sensing distance seemed to be an important value that was missing from the literature. In this paper, we model the radiation transfer process from a 37 degrees C object, i.e. warm-blooded prey, to an infrared pit organ. The model tries to answer a very basic question-at what distance does the thermal signature of a 37 degrees C object blend into the background for a non-imaging biological infrared sensor? The output of the model, the sensing distance, is of particular interest in comparing biological infrared sensors to current semiconductor-based infrared (IR) detectors-largely because of inappropriate comparisons between the temperature sensitivity of IR snake reception and imaging IR cameras. The purpose of the presented work to make more appropriate comparisons, i.e. sensing distance. This sensing distance output indicates an extremely short detection distance (<5 cm)-contradictory to what is observed experimentally. This dichotomy raises further questions regarding how the biological system amplifies this weak signal.     

Surface ultrastructure of pit organ, spectacle, and non pit organ epidermis of infrared imaging boid snakes: A scanning probe and scanning electron microscopy study.

Boid snakes possess unique infrared imaging pit organs. The ultrastructure of the surfaces of these organs scatter or reflect electromagnetic radiation of specific wavelengths. Pit organ epidermal surfaces of boid snakes are covered with arrays of pore-like structures called micropits. In order to determine the dimensions of this complicated surface structure, we have performed the first ultrastructural analysis on snake epidermis by high-resolution microscopy techniques. Using scanning probe microscopy and scanning electron microscopy, we found that the epidermis of pit organ, maxillary non pit organ, spectacle, and ventral scales contain arrays of micropits. These scale surfaces also contain major surface features of overlapping plate-like structures. Pit organ micropits averaged 319 nm in diameter and 46 nm in depth and were spaced an average of 808 nm from each other. These micropits were significantly deeper, of greater diameter, and spaced at greater distances apart than those of the other scales. Plate structures of the pit organs had a mean distance between plates of 3.5 microm and a mean plate step height of 151 nm. These differences serve to strengthen the argument that arrays of micropit and plate surface structures function as spectral filters or anti-reflective coatings with respect to incident electromagnetic radiation.     

Responses of the infrared sensilla of Melanophila acuminata (Coleoptera: Buprestidae) to monochromatic infrared stimulation

The pit organs of the beetle Melanophilaacuminata were stimulated with monochromatic infrared radiation using a continuous wave CO overtone infrared laser. Best sensitivity was in the wavelength range 2.8–3.5 μm. In this range a stimulus intensity of 14.7 mW cm⁻² was sufficient to generate single action potentials. At a wavelength of 5 μm receptor performance significantly decreased. An increase in stimulus intensity caused a decrease in response latency and an increase in the number of action potentials elicited. At a given wavelength (3.4 μm) the dynamic amplitude range of action potential responses covered 12 dB. At high stimulus intensities (94.2 mW cm⁻²) a stimulus duration of 4 ms was sufficient to generate one to two action potentials and a stimulus duration of 60 ms already caused response saturation (with up to nine action potentials). In a repetitive stimulus regime distinct receptor potentials were visible up to a frequency of 600 Hz. Accepted: 18 March 2000     

Functional analysis of actin fibrils in Physarum polycephalum

Summary Fluorochromed heavy meromyosin (TRITC-HMM) was microinjected as a molecular probe into small sandwich-plasmodia of Physarum polycephalum with the aim to demonstrate the spatial morphology and to analyze the dynamic activity of the fibrillar actin system in the living state. The plasmodia display different fibrillar organizations with a polygonal arrangement in the front region (FR) and a parallel or helical arrangement along protoplasmic veins in the intermediate (IR) and uroid region (UR). Quantitative evaluations by measuring the total length, lifetime, dynamic activity, long-term stability and optical density of fibrils reveal distinct differences between the three plasmodial regions: The total length (FR = 27.1 ± 18.5 m, IR = 24.8 ± 12.9 m, UR= 12.3 ± 4.7 m), the lifetime (FR = 12.2 ± 3.4 min, IR=10.5 ± 3.7 min, UR = 6.0 ± 3.4 min), and the dynamic activity as measured in length changes per min (FR = 17.9 ± 11.3 m, IR = 13.1 ± 3.9 m, UR = 8.3 ± 3.9 m) distinctly decrease from the front to the uroid region. On the other hand, the greatest stability as determined by lifetime changes in length (FR = -2.4 ± 16.2 m, IR = 0.3 ± 10.1 m, UR = -6.6 ± 8.9 m) and the highest optical density as expressed in grey-values (FR = 57.0 ± 14.1 gv, IR = 115.6 ± 26.1 gv, UR 62.5 ± 8.1 gv) were found for actomyosin fibrils of the intermediate region. The morphological and physiological data of the present paper are discussed with respect to the biological significance of the fibrillar microfilament system in Physarum polycephalum.     

Intracellular optical physiology of the bee's eye

Summary Intracellular optical physiology is a newly developed, non-invasive technique for recording from single types of insect photoreceptor cells. An intact animal is mounted on the goniometer stage of a double-beam, incident-light microspectrophotometer. The stimulating beam delivers monochromatic flashes that evoke pupillary responses from photoreceptor cells in a localized region of the eye. The measuring beam delivers red or infrared illumination that continuously measures the changes in reflectance that accompany the pupillary response. In this paper on the worker bee we demonstrate two experimental conditions under which only one of the three spectral types of photoreceptor dominates the measured response.When the eye is dark-adapted, and increases in reflectance are measured with an infrared beam, threshold responses evoked by long flashes (40 s or more) are dominated by that cell type which is most sensitive to the stimulus. The following data were taken from the dorsal (ventral) poles of the bee's eye: At 350 nm, a quantum flux of 0.1 (0.05)×10 ¹² photons/cm ²/s/numerical aperture of 0.2 produces a threshold response that originates only from the UV-receptors. At 430 nm, a quantum flux of 0.6 (0.2) units produces a threshold response that is dominated by the blue-receptors. At 530 nm, a flux of 1.0 (0.1) units produces a threshold response that originates only from the green-receptors.When the eye is red-adapted, the pupillary responses to short flashes (10 s or less) are dominated by the green-receptors. Under this condition the pupillary action-spectrum for sensitized green-receptors is the same as the electrophysiological spectral sensitivity function of the minimally coupled green-receptor. This is true throughout the spectral range 350 nm–650 nm (Fig. 3).If the duration of stimulating flashes is increased to 40 s, the shape of the action spectrum is unchanged for wavelengths greater than 470 nm, but is significantly elevated, by as much as 1.8 log-units, for wave-lengths less than 420 nm (Fig. 4). In this case the UV-receptors dominate the pupillary response at short wavelengths, while the green-receptors dominate it at long wavelengths.We used these effects to determine that all three spectral types of receptor are present in the regions of both dorsal and ventral poles, as well as in the frontal region of the eye.This work was supported by grants EY01140 and EY00785 from the National Eye Institute, U.S.P.H.S. (to GDB), by the Connecticut Lions Eye Research Foundation (to GDB), and by a grant from the University of Zurich (to RW), and by grant 3.529.075 from the Swiss National Science Foundation (to RW) including a Senior Research Fellowship awarded to GDB. We thank Dr. Thomas Labhart for many helpful discussions and for allowing us to refer to his unpublished data, and Dr. Doekele Stavenga for his critical, constructive comments.     

Thresholds of retinal and extraretinal photoreceptors measured by photobehavioral response in catfish,Silurus asotus

Summary Light thresholds of retinal and extraretinal photoreceptors in catfish were examined by the photobehavioral response using a method in which reflex body movements are recorded. Thresholds were determined in six groups, (A) intact, (B) ophthalmectomized, (C) pinealectomized, (D) ophthalmectomized + pinealectomized, (E) ophthalmectomized, pinealectomized and skinless over the brain (skinless fish) and (F) ophthalmectomized, pinealectomized and dorsally covered with aluminum foil over the brain (covered fish). All these fishes displayed short term activity to white light stimulation after being dark adapted for more than 5 h. The lowest threshold was obtained in the intact group (2.0×10^–4 W/cm²). The thresholds of lateral eyes and the pineal organ were 3.4×10^–3 and 1.5×10^–2 W/cm², respectively. Without lateral eyes and pineal organ, catfish still responded to light, indicating the possible existence of extraretinal nonpineal photoreceptors (ENPs). The threshold of ENPs was 3.3 W/cm². The localization of ENPs was assumed to be in the brain from the experiment with the combination of skinless and covered fish.Abbreviation ENP extraretinal nonpineal photoreceptor     

A spectral analysis of stem bark for boreal and temperate tree species

The woody material of forest canopies has a significant effect on the total forest reflectance and on the interpretation of remotely sensed data, yet research on the spectral properties of bark has been limited. We developed a novel measurement setup for acquiring stem bark reflectance spectra in field conditions, using a mobile hyperspectral camera. The setup was used for stem bark reflectance measurements of ten boreal and temperate tree species in the visible (VIS) to near‐infrared (NIR) (400–1000 nm) wavelength region. Twenty trees of each species were measured, constituting a total of 200 hyperspectral reflectance images. The mean bark spectra of species were similar in the VIS region, and the interspecific variation was largest in the NIR region. The intraspecific variation of bark spectra was high for all studied species from the VIS to the NIR region. The spectral similarity of our study species did not correspond to the general phylogenetic lineages. The hyperspectral reflectance images revealed that the distributions of per‐pixel reflectance values within images were species‐specific. The spectral library collected in this study contributes toward building a comprehensive understanding of the spectral diversity of forests needed not only in remote sensing applications but also in, for example, biodiversity or land surface modeling studies.     

Reduced performance of prey targeting in pit vipers with contralaterally occluded infrared and visual senses

Both visual and infrared (IR) senses are utilized in prey targeting by pit vipers. Visual and IR inputs project to the contralateral optic tectum where they activate both multimodal and bimodal neurons. A series of ocular and pit organ occlusion experiments using the short-tailed pit viper (Gloydius brevicaudus) were conducted to investigate the role of visual and IR information during prey targeting. Compared with unoccluded controls, snakes with either both eyes or pit organs occluded performed more poorly in hunting prey although such subjects still captured prey on 75% of trials. Subjects with one eye and one pit occluded on the same side of the face performed as well as those with bilateral occlusion although these subjects showed a significant targeting angle bias toward the unoccluded side. Performance was significantly poorer when only a single eye or pit was available. Interestingly, when one eye and one pit organ were occluded on opposite sides of the face, performance was poorest, the snakes striking prey on no more than half the trials. These results indicate that, visual and infrared information are both effective in prey targeting in this species, although interference between the two modalities occurs if visual and IR information is restricted to opposite sides of the brain.     

粘虫下唇须陷窝器及其感器的形态、类型与分布

【目的】明确我国重要农业害虫粘虫Mythimna separata Walker下唇须和陷窝器形态结构以及陷窝器内部感器的形态、类型与分布。【方法】利用光学显微镜观察粘虫成虫下唇须及陷窝器形态结构,利用扫描电子显微镜观察陷窝器内部的感器。【结果】结果表明,粘虫成虫下唇须似管状,具3节,各节形态和长度不同,其中第2节最长。粘虫下唇须长度雌雄异形,雌性的下唇须长度为2 463.50±143.65 μm,显著短于雄性的（2 566.11±70.28 μm）（t=2.722, df=34, P=0.012）。陷窝器凹坑深约280 μm,开口处直径约50 μm,内部直径约32 μm,雌雄间无显著差异。陷窝器内部的感器主要包括毛形感器和棒状感器2种类型。毛形感器位于陷窝器凹坑的上半部分,而棒状感器分布在陷窝器凹坑的下半部分。雌性毛形感器长为18.20±0.84 μm,显著短于雄性的21.24±0.34 μm（t=3.335, df=30, P=0.003）。而雌性的棒状感器长为14.69±0.48 μm,显著长于雄性的12.31±0.49 μm（t=3.452, df=21, P=0.002）。【结论】粘虫下唇须陷窝器感器分属2大类型,分布于陷窝器内不同的区域,其长度具有性别差异性。本文报道的这些形态学观察结果为进一步研究粘虫下唇须陷窝器感器生理和功能奠定了基础。     

Frequency response of the electroreceptors (“small pit organs”) of the catfish,Ictalurus nebulosus LeS

Summary By electrophysiologically recording the spikes from the ramus lateralis ventralis nervi vagi of the catfishIctalurus nebulosus LeS., we determined the frequency response of the electroreceptors (small pit organs) in unanaesthetized, freely swimming specimens. The response frequency domain we were able to measure ranged from 0,03 to 25 Hz. We found a peak of greatest sensitivity at about 3 to 7 Hz (Fi. 5), using a stimulus with a current density of 5 × 10^–4A/mm² in water with a specific resistance of 20 Ohm. m. The threshold current density of the small pit organs in a homogeneous sinusoidally varying electric field (3 Hz) proved to be 3 × 10^–5 A/mm².

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Zusammenfassung Durch Ableitung von Nervenimpulsen vom ramus lateralis ventralis nervi vagi beim Zwergwels,Ictalurus nebulosus LeS., versuchten wir den Frequenzbereich der Electroreceptoren (small pit organs) bei frei herumschwimmenden, nicht narkotisierten Tieren zu bestimmen. Der wirksame Frequenzbereich, in dem Bestimmungen vorgenommen werden konnten, erstreckte sich von 0,03 bis 25 Hz. Die Empfindlichkeitskurve der small pit organs erreichte einen Gipfel bei Frequenzen von 3–7 Hz (Fig. 5). Die Reizstromdichte betrug dabei 5 × 10^–4A/mm²; der spezifische Wasserwiderstand war 20 Ohm. m. Es stellte sich heraus, daß die Schwellenstromdichte der small pit organs im homogenen elektrischen Wechselstromfeld (3 Hz) 3 × 10^–5 A/mm² betrug.

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We wish to thank Prof. S. Dijkgraaf for his stimulating interest, and Mr. W. J. G. Loos for making the drawings.     

Effect of methoprene and 20-hydroxyecdysone on vitellogenin production in cultured fat bodies and backless explants from unfed female Dermacentor variabilis

The effect of 20-hydroxyecdysone (20HE) and the juvenile hormone analogue methoprene (JHA) on vitellogenin (Vg) production in fat body organ cultures and backless explants of unfed female Dermacentor variabilis was measured. An indirect double antibody enzyme linked immunosorbent assay (ELISA) was developed using a monoclonal antibody that recognized a 98 kDa subunit of Vg and a Vg specific polyclonal antibody made against vitellin (Vn). Peak Vg titers in culture medium from fat body cultures treated with 0.1 &mgr;M 20HE or 1 &mgr;M 20HE were 24 ng/ml and 20 ng/ml respectively. In culture medium from backless explants treated with 0.1 &mgr;M 20HE or 1 &mgr;M 20HE, peak Vg titers were 36 ng/ml and 26 ng/ml, respectively. JHA produced only a slight increase in Vg titers that was statistically different from Vg titers produced by 20HE but was not statistically different from hormone-free controls. These results support the conclusion that Vg production in fat body trophocytes of D. variabilis is regulated by 20HE.     

Endocytosis and membrane turnover in the germ tube of uromyces fabae

We have used the fluorescent dye FM4-64 as a tracer to demonstrate bulk membrane internalization (endocytosis) and redistribution of the dye within the cytoplasm of the germ tube of the rust fungus Uromyces fabae. Staining of the hyphal membrane was detected 4 s after application of FM4-64 and reached a maximum after 1 min. The highest fluorescence intensity occurred in the apex. Subsequently, staining of the plasma membrane decreased and a subapical region of the fungal protoplast (5-20 &mgr;m from the tip) displayed increasing fluorescence with a maximum after 5 min. Fluorescence in the subapical region was redistributed to an area in the hyphal tip, which corresponds to the accumulation of apical vesicles, after 10-15 min and subsequently to a cytoplasmic region in front of the two nuclei (35-45 &mgr;m from the tip). We conclude from our measurements of membrane fluorescence that the turnover time from endocytosis to secretion of the dye amounts to 15 min. The uptake of the dye into the cytoplasm, but not membrane loading, could be inhibited completely with 5 mM NaN3 or by a temperature shift to 4 degreesC. This is the first evidence for endocytosis in a fungal germ tube. Copyright 1998 Academic Press.     

Maturation of optics and resolution in adult dung beetle superposition eyes

Summary The maturation of the morphology and refractive-index gradients of the crystalline cones and cornea of the superposition compound eye of the nocturnal dung beetle Onitis aygulus was traced as a function of age following adult ecdysis. Intracellular recordings from retinula cells were also made to trace the maturation of angular sensitivity, thereby determining the resolution of the eye at each age. Maturation proceeded quickly during the first 3 days following ecdysis, and then more slowly, with full maturation attained by the end of the second week. The main experimental results obtained during the first 2 weeks after ecdysis were: (1) the mean length of the crystalline cones increases from 67 m to 79 m; (2) the mean thickness of the cornea increases from 36 m to 50 m; (3) the mean refractive index along the axis of the crystalline cones increases from 1.459 to 1.511 in the distal cone region, from 1.434 to 1.501 in the waist region and from 1.425 to 1.486 in the proximal region; (4) the shape of the refractive-index gradient becomes more parabolic; (5) the mean rhabdom cross-sectional area increases from 190 m² to 230 m²; (6) the angular-sensitivity function narrows, with the acceptance angle decreasing from 22 ° to 4 °. An optical ray tracing model predicts an image quality limited only by the immaturity of the optics, a prediction confirmed by the electrophysiological results. The results are compared to optical maturation in day-active moths and skipper butterflies (which mature very quickly) and discussed in relation to ecological efficiency.