Visual afferences to flight steering muscles controlling optomotor responses of the fly

Summary In tethered flying house-flies (Musca domestica) visually induced turning reactions were monitored under open-loop conditions simultaneously with the spike activity of four types of steering muscles (M.b1, M.b2, M.I1, M.III1). Specific behavioral response components are attributed to the activity of particular muscles. Compensatory optomotor turning reactions to large-field image displacements mainly occur when the stimulus pattern oscillates at low frequencies. In contrast, turning responses towards objects are preferentially induced by motion of relatively small stimuli at high oscillation frequencies. The different steering muscles seem to be functionally specialized in that they contribute to the control of these behavioral responses in different ways. The muscles I1, III1 and b2 are preferentially active during small-field motion at high oscillation frequencies. They are much less active during small-field motion at low oscillation frequencies and large-field motion at all oscillation frequencies which were tested. M.b2 is most extreme in this respect. These steering muscles thus mediate mainly turns towards objects. In contrast, M.b1 responds best during large-field motion at low oscillation frequencies and, thus, is appropriate to control compensatory optomotor responses. However, the activity of this muscle is also strongly modulated during small-field motion at high oscillation frequencies and, therefore, may be involved also in the control of turns towards objects. These functional specializations of the different steering muscles in mediating different behavioral response components are related to the properties of two parallel visual pathways that are selectively tuned to large-field and small-field motion, respectively.Abbreviations FD (cell) figure detection (cell) - HS (cell) horizontal (cell)     

Dynamic properties of two control systems underlying visually guided turning in house-flies

Summary The compensatory optomotor turning reaction as well as the turning response towards objects play an important role in visual orientation. On the basis of behavioural experiments under precisely defined stimulus conditions it is concluded that in female house-flies these motion-dependent responses are mediated by two parallel control systems with different dynamic and spatial integration properties. One of them (large-field system) is most sensitive to the motion of large textured patterns and controls the yaw torque mainly at low oscillation frequencies (below 0.1 Hz) of the stimulus panorama. In contrast, the other control system (small-field system) is tuned to the detection of relatively small moving patterns and shows its strongest responses at high oscillation frequencies (between 1 and 4 Hz), i.e. in a frequency range where the large-field system contributes to the turning response with only a relatively small gain.In free flight, house-flies do not curve smoothly but in sequences of rapid turns which induce retinal large-field motion of continually changing sign (Wagner 1986b). The dynamic properties of the large-field system might thus be interpreted as a simple strategy to almost eliminate the unwanted optomotor yaw torque induced by active self-motion. In contrast, the small-field system might still be operational under these conditions.     

Elementary detectors for vertical movement in the visual system of Drosophila

Optomotor thrust responses of the fruitfly Drosophila melanogaster to moving gratings have been analysed in order to determine the arrangement of elementary movement detectors in the hexagonal array of the compound eye. These detectors enable the fly to perceive vertical movement. The results indicate that, under photopic stimulation of a lateral equatorial eye region, the movement specific response originates predominantly from two types of elementary movement detectors which connect neighbouring visual elements in the compound eye. One of the detectors is oriented vertically, the other detector deviates 60° towards the anterior-superior direction (Fig. 5b). The maximum of the thrust differences to antagonistic movement is obtained if the pattern is moving vertically or along a superior/anterior — inferior/posterior direction 30° displaced from the vertical (Fig. 3d,e, Fig. 6). Only one of the detectors coincides with one of the two detectors responsible for horizontal movement detection. This indicates that a third movement specific interaction in the compound eye of Drosophila has to be postulated. — The contrast dependence of the thrust response (Fig. 2) yields the acceptance angle of the receptors mediating the response. The result coincides with the acceptance angle found by analysis of the turning response of Drosophila (Heisenberg and Buchner, 1977). This value corresponds to the acceptance angle expected, on the basis of optical considerations, for the receptor system R 1–6. — The movement-specific neuronal network responsible for thrust control is not homogeneous throughout the visual field of Drosophila. Magnitude and preferred direction of the thrust response in the upper frontal part of the visual field seem to vary considerably in different flies (Fig. 6).     

Orientation sensitivity of the visual movement detection system activating the landing response of the blowflies,Calliphora, and Phaenicia: A behavioural investigation

The orientation sensitivity of the visual movement detection system relative to the axes of the eye was investigated for the landing response by changing the direction of movement of a periodic striped pattern (unidirectional movement) and a two-stripe pattern consiting of two stripes moving apart (bidirectional movement). In the momocular, equatorial regions of the eye progressive motion proves to be most effective, whereas in the frontal (equational), binocular region descendive motion is most effective in eliciting the landing response, probably caused by binocular interactions. A strong enhancement of the response is induced by stimulation in the binocular region of the two eyes. The orientation of elementary movement detectors relative to the axes of the ommatidial array is discussed. The findings are summarized in a functional model of the landing response.     

Characteristics of Gamma-Oscillations in Responses of Neurons of the Cat Lateral Geniculate Body

There were studied characteristics of gamma-oscillations in responses of neurons of the lateral geniculate body (LGB) in cat to exposure in their receptive fields (RF) of half-tone and binary test images. The gamma-oscillations were observed in 38.8% of cases (69 cells). The spectral characteristics (SC) (the band 20–100 Hz) of the neuronal responses to adequate stimuli (on- and off-responses correspondingly of on- and off-neurons) were analyzed. The total of 5930 poststimulus histograms (PSTH) of responses constructed from 177 900 neuronal impulse responses were considered. The mean value of the SC dominant frequencies of the whole sample of the neuronal responses amounted to 44.74 ± 21.46 Hz. In this cell sample, the neurons were revealed, which generated oscillations with markedly different frequencies in response to the same stimuli. Based on this property, three types of neurons were determined, with the mean oscillation frequencies of 26.95 ± 4.35, 52.02 ± 9.05, and 85.79 ± 7.19 Hz. The histograms of distribution of peak frequency values in SC of the neuronal responses and of index values of these oscillation peaks also revealed three maxima that corresponded to the frequencies of the three described types of neurons. The mean values of dominant frequencies of gamma-oscillations in responses of all three types of neurons remained constant (within the limits of dispersion) at changes of spatial-brightness parameters of test stimuli as well as at changes of the neuronal excitation level (the number of impulses in responses). The oscillation index values of dominant frequencies depended on parameters of the test images and correlated with the neuronal excitation level (the coefficient of correlation was 0.78 from data of 5930 CX). The suggestion is made about the existence in the neuronal network of the synchronization mechanisms functioning on the principle of multiple synchronization.      

Die optischen Übertragungseigenschaften der Komplexaugen von Drosophila

Summary The transfer properties of the optical system in the arthropod compound eye are determined by the interommatidial angle , influencing the resolving power, and by the width of the visual fields of single ommatidia , influencing the response at high spatial frequencies of brightness distributions in the object space. The energy transfer/ receptor is proportional to ( )² and decreases with in-inreasing approximation of the perfect-imaging condition: gD 0; 0. However, a value > 0 has to be maintained in order to overcome the threshold of nervous excitation at a certain minimum-brightness level. Theoretical treatment yields /=0.62 to 0.88 as the corresponding optimum-imaging relation. The actual ratio can be derived from measurements of the optomotor reactions to the movement of periodic brightness patterns. The approximate value 0.76 is obtained from the fruitfly Drosophila with normal and mutant eye pigmentation. As a result, the parameters of this imaging system are found to be established in a way that enables optimum performance at sufficient illumination. An dieser Stelle möchte ich Dr. W. Reichardt für sein eingehendes Interesse und manche anregende Diskussion über die Sehvorgänge im Komplexauge meinen Dank sagen. Dr. K. Kirschfeld verdanke ich ebenfalls wertvolle Hinweise. Herrn E. Freiberg bin ich für die Anfertigung der Abbildungen sehr verbunden.     

Elementary movement detectors in an insect visual system

In the theoretical part of the present work the input-output relation for a multi-input system is developed into a functional power series. This is formally equivalent to a decomposition of the system into a sum of all possible combinations of 1-, 2-, 3-... input subsystems. The average response of the system to a uniformly moving patern is known to be a Fourier series with respect to spatial frequency. The coefficients of the series are linear combinations of the weights by which different subsystems contribute to the total reaction. If a system can be shown to have essential nonlinearities of no higher than second order it is possible to calculate, from a Fourier analysis of the average movement response, the weight by which the nonlinear interaction between any two input elements contributes to the total reaction. This interaction is termed elementary movement detector. By the analysis presented here the arrangement of the elementary movement detectors may be determined for a two-dimensional array of input elements and the strength of their contributions to the total movement reaction may be calculated. Special experimental methods have been developed which allow one to apply this analysis to the visual system of the fruitfly Drosophila. The preliminary data presented show that the direction sensitive optomotor response of Drosophila can be attributed predominantly to the contributions from two elementary movement detectors which interconnect neighbouring visual elements. The detectors are oriented in the hexagonal array of the compound eye at +30° and at-30° with respect to the horizontal line of symmetry. A weak contribution from a detector between neighbouring elements along the horizontal line of symmetry is suggested by the present data. In the course of the analysis the contrast transfer properties of the compound eye are characterized.     

The function and role of two types of mechanoreceptive ‘free’ nerve endings in the head skin of amphibian embryos

Summary The properties of trigeminal ganglion sensory neurones innervating the head skin of lateXenopus laevis embryos have been studied using extracellular recordings. Two types of mechanosensory neurones were found:Rapid-transient detectors which responded with few impulses to rapid, local, indentation of the skin with a fine probe (10–25 m diameter), andMovement detectors which responded with a slowly adapting discharge to even very slow distortion of the skin (5 m· s^–1) with small or large probes. Receptive fields over the whole head surface as far back as the gill rudiments were plotted for both types of neurone. The areas for the two types were similar (means of 0.015 mm² for rapid-transient and 0.017 mm² for movement).Comparative observations on embryos ofRana temporaria andTriturus helveticus showed a very similar division of trigeminal sensory neurones into two types. InXenopus embryos stimuli which only excite the movement detectors were found to have inhibitory effects on behaviour. They would stop swimming and responses to other excitatory stimuli. Stimuli which excited the rapid-transient detectors normally evoked swimming. The division of the somatosensory system inXenopus embryos into two subsystems with different sensitivities and inhibitory, or excitatory effects on behaviour is discussed and related to findings in other groups of animals.I thank the MRC for support.     

Solid state fermentation of straw withNeurospora crassa for CMCase and β-glucosidase production

Summary CMCase and -glucosidase were produced by the mutantNeurospora crassa 40b cultivated on untreated wheat straw in a solid state fermentation. Best enzyme activities were observed when the growth medium was composed of wheat straw mixed with certain mineral solutions at a ratio 1:2 (w/v). A partially purified enzyme preparation showed optimum enzyme activities of CMCase and -glucosidase at pH 4.0 and 5.0 and temperature 50 and 60°C respectively. The apparent Km values for the same enzymes were 16.8 g/l and 1.03x10^–4 M respectively. At optimum growth and enzyme assay conditions yields as high as 586.2 U CMCase and 58.4 U -glucosidase per gram of straw were obtained.     

Role of bag cells in egg deposition ofAplysia brasiliana

Summary Egg deposition behaviors are analyzed from time-lapse recordings during which spontaneous discharges of the neuroendocrine bag cells are recorded with chronically implanted cuff electrodes. In the laboratory,Aplysia brasiliana normally deposit long egg cordons on the substrate in a characteristic figure 8 pattern similar to the configuration of egg masses observed in the natural environment. The overt behaviors associated with egg deposition are rhythmic head movements consisting of three components that overlap with characteristic relative latencies: up-and-down undulations, side-to-side weaves and in-and-out tamps. The characteristics of the three behaviors and their time courses relative to the appearance of eggs on the substrate suggest that undulations prepare the substrate, weaves distribute the egg cordon and tamps attach the cordon to the substrate. The same rhythmic head movements are also elicited by injections of homogenized abdominal ganglia (HAG) containing bag cell clusters, with comparable relative latencies and maximum frequencies but for shorter total durations. The overt behaviors begin earlier for normal than for triggered egg laying, often before the spontaneous release of bag cell hormones. This suggests that the head oscillations in intact animals are not normally initiated by bag cell activity. The mean latency to the appearance of the egg cordon on the substrate is the same (about 34 min) following either HAG injections or spontaneous bag cell discharges, confirming previous suggestions that the bag cell discharge triggers ovulation. Furthermore, the head movements appear to terminate at the same time following release or injection of hormone. The accompanying paper demonstrates that the full expression of the behavioral effects of bag cell injections depend upon normal movement of eggs in the reproductive tract.Abbreviations CPG central pattern generator - ELH egg laying hormone - HAG homogenized abdominal ganglia     

Electromotility of outer hair cells from the cochlea of the echolocating bat,Carollia perspicillata

Isolated outer hair cells (OHCs) and explants ot the organ of Corti were obtained from the cochlea of the echolocating bat, Carollia perspicillata, whose hearing range extends up to about 100 kHz. The OHCs were about 10–30 m long and produced resting potentials between-30 to -69 mV. During stimulation with a sinusoidal extracellular voltage field (voltage gradient of 2 mV/m) cyclic length changes were observed in isolated OHCs. The displacements were most prominent at the level of the cell nucleus and the cuticular plate. In the organ of Corti explants, the extracellular electric field induced a radial movement of the cuticular plate which was observed using video subtraction and photodiode techniques. Maximum displacements of about 0.3–0.8 m were elicited by stimulus frequencies below 100 Hz. The displacement amplitude decreased towards the noise level of about 10–30 nm for stimulus frequencies between 100–500 Hz, both in apical and basal explants. This compares well with data from the guinea pig, where OHC motility induced by extracellular electrical stimulation exhibits a low pass characteristic with a corner frequency below 1 kHz. The data indicate that fast OHC movements presumably are quite small at ultrasonic frequencies and it remains to be solved how they participate in amplifying and sharpening cochlear responses in vivo.Abbreviations BM basilar membrane - FFT fast Fourier Transfer - IHC inner hair cell - OHC outer hair cell     

Dynamics of pattern formation

Experiments were designed to test the validity ofTuring's suggested pattern-forming mechanisms, which are initially capable of giving rise to only five to seven uniform structures.TheOregon-R (wild-type), mass-cultured strain ofDrosophila melanogaster was employed. Selection for the regular arrangement of microchaetac on the margin of the fourth abdominal sternite was practiced for twenty generations. In the L line, individuals with six uniformly spaced bristles were selected as parents of every generation. Due to the absence of nine bristles dividing the sternal margin uniformly, the progeny was raised in each generation in the H line from males and females with nine as equidistant bristles as possible. The whole experiment was performed at 25±0.50°C.Selection was effective in increasing the frequency of six regular bristles in the L line. But no progress in the desired direction was obtained in the H line, although the proportion of sternites with nine irregular structures was found to increase. The experimental results supportTuring's diffusion-reaction scheme of pattern formation in morphogenesis.Supported by grants GB-1388 and GB-3219 from the National Science Foundation.     

Meiotic studies ofElymus nutans andE. jacquemontii (Poaceae,Triticeae) and their hybrids withPseudoroegneria spicata and seventeenElymus species

Hybridizations ofElymus nutans andE. jacquemontii were carried out with one species ofPseudoroegneria (S genome), and 20Elymus species, each containing either of the SH, SY, SYH, or SYW genomes. Chromosome configurations were analysed at metaphase I of the two target taxa and their interspecific hybrids. It is concluded that (i)E. nutans is an allohexaploid containing the SYH genomes, andE. jacquemontii is an allotetraploid having the SY genomes; (ii) the genomic affinity is associated with the geographic distance between the species studied; (iii) minor genomic structural rearrangements have occurred within the hexaploid taxon ofE. nutans.     

Group activity and population dynamics of the howler monkey on Barro Colorado Island

The mantled howler monkey (Alouatta palliata) of Barro Colorado Island, Panama Canal Zone, was studied in the late rainy season of 1970. Group ranges, group activity patterns and population size and structure were recorded for six groups in a 204 hectare study area. A new trail survey method was introduced to determine the population size of the entire island. Using this method, a population of 923±125 animals is estimated. Factors affecting group stability and group density are discussed. Rainfall appears to be a major determinant of howler activity and group movement in the late rainy season.     

An elaborated model of fly small-target tracking

Flies have the capability to visually track small moving targets, even across cluttered backgrounds. Previous computational models, based on figure detection (FD) cells identified in the fly, have suggested how this may be accomplished at a neuronal level based on information about relative motion between the target and the background. We experimented with the use of this small-field system model for the tracking of small moving targets by a simulated fly in a cluttered environment and discovered some functional limitations. As a result of these experiments, we propose elaborations of the original small-field system model to support stronger effects of background motion on small-field responses, proper accounting for more complex optical flow fields, and more direct guidance toward the target. We show that the elaborated model achieves much better tracking performance than the original model in complex visual environments and discuss the biological implications of our elaborations. The elaborated model may help to explain recent electrophysiological data on FD cells that seem to contradict the original model.Acknowledgement This work was supported by the US Office of Naval Research under agreement number N68936-00-2-0002.     

Conformational Change of the Rieske [2Fe–2S] Protein inCytochrome bc 1 Complex

Structures of mitochondrial bc ₁ complex have been reported based on four different crystalforms by three different groups. In these structures, the extrinsic domain of the Rieske [2Fe–2S]protein, surprisingly, appeared at three different positions: the c ₁ position, where the [2Fe–2S]cluster exists in close proximity to the heme c ₁; the b position, where the [2Fe–2S] clusterexist in close proximity to the cytochrome b; and the intermediate position where the[2Fe–2S] cluster exists in between c ₁ and b positions. The conformational changes betweenthese three positions can be explained by a combination of two rotations; (1) a rotation of theentire extrinsic domain and (2) a relative rotation between the cluster-binding fold and thebase fold within the extrinsic domain. The hydroquinone oxidation and the electron bifurcationmechanism at the Q_P binding pocket of the bc ₁ complex is well explained using theseconformational changes of the Rieske [2Fe–2S] protein.     

Enzymatic β-galactosidation of β-xylopyranosides

Summary The disaccharides formed by enzymatic transfer of the -D-galactopyranosyl residue fromo-nitrophenyl -d-galactopyranoside to -d-xylopyranosides have been identified. The influence of different factors on the yields of the disaccharides obtained was evaluated. Significant changes in selectivity were observed when -galactosidase fromE. coli was used instead of -galactosidase fromA. oryzae.     

Memorization and delayed expression of regulatory messages in plants

Plantlets of Bidens pilosus L., considered to be basically symmetrical, can be lateralized (A/B) by being administered a symmetry-breaking signal such as puncturing one of the plant cotyledons. The induced asymmetry remains latent as long as the plants have not been made permissive, i.e. as long as the plant apex is left functioning. When the apex has been removed (plant decapitation), the latent asymmetry is expressed by one of the cotyledonary buds (a/b) statistically beginning to elongate before the other. The interval of time between delivering the symmetry-breaking signal and making the plant permissive is the memorization-time, t. Memorization can be quantified by using a precedence index, q, the values of which range from 0 (no detectable asymmetry with regard to bud growth) to ±1 (bud growth perfectly asymmetric in favour of either bud b or a). Even for memorization times, t, up to 14 d, q-values up to 0.4 (or even larger) are observed. Various experimental characteristics (e.g. light, temperature, presence or absence of the root system) but not the plant age can affect the q-values, at the moment when the treatments are performed, at least in the range of 6 to 25 d. Combining several puncturing treatments either increases or decreases the q-values, depending on the nature of these treatments and the time-intervals, t, between them. Symmetrically removing both cotyledons in the minutes following the puncturing of one of them does not significantly alter the results, which means that the symmetry-breaking message is rapidly transported and memorized within the plant. Non-traumatic asymmetrical treatments (droplets of saline solutions, light-gradients) can also act as symmetry-breaking signals and be memorized. Plants other than Bidens are likely to possess similar memorization ability, although the q-values observed up to now have not been very large.     

The Effect of Salicylic Acid on Peroxidase Activity in Wheat Calli Cocultured with the Bunt Pathogen Tilletia caries

The effect of salicylic acid (SA) on peroxidase activity in wheat (Triticum aestivum L.) calli cocultured with the bunt pathogen Tilletia caries was studied. Fungal infection was shown to activate cytoplasmic peroxidase. SA suppressed total peroxidase activity but did not inhibit the peroxidase with pI 9.8. A novel chitin-specific peroxidase with pI 3.5 appeared after the SA treatment. The infection of SA-treated cells with Tilletia caries activated the isoenzymes with pI 3.5, 4.8, and 7.5 and stimulated their secretion into the culture medium. The ability of SA to control wheat peroxidase activity during pathogenesis is discussed. The important role of this control in plant defense responses to the bunt pathogen is emphasized.     

Visual detection of paradoxical motion in flies

Summary From psychophysics it is known that humans easily perceive motion in Fourier-stimuli in which dots are displaced coherently into one direction. Furthermore, motion can be extracted from Drift-balanced stimuli in which the dots on average have no distinct direction of motion, or even in paradox -motion stimuli where the dots are displaced opposite to the perceived direction of motion. Whereas Fourier-motion can be explained by very basic motion detectors and nonlinear preprocessing of the input can account for the detection of Drift-balanced motion, a hierarchical model with two layers of motion detectors was proposed to explain the perception of -motion. The well described visual system of the fly allows to investigate whether these complex motion stimuli can be detected in a comparatively simple brain.The detection of such motion stimuli was analyzed for various random-dot cinematograms with extracellular recordings from the motion-sensitive Hl-neuron in the third visual ganglion of the blowfly Calliphora erythrocephala. The results were compared to computer-simulations of a hierarchical model of motion detector networks.For Fourier- and Drift-balanced motion stimuli, the Hl-neuron responds directionally selective to the moving object, whereas for -motion stimuli, the preferred direction is given by the dot displacement. Assuming nonlinear preprocessing of the detector input, such as a half-wave rectification, elementary motion detectors of the correlation type can account for these results.Abbreviations EMD elementary motion detector