The three-dimensional optomotor torque system ofDrosophila melanogaster

Summary The well known optomotor yaw torque response in flies is part of a 3-dimensional system. Optomotor responses around the longitudinal and transversal body axes (roll and pitch) with strinkingly similar properties to the optomotor yaw response are described here forDrosophila melanogaster. Stimulated by visual motion from a striped drum rotating around an axis aligned with the measuring axis, a fly responds with torque of the same polarity as that of the rotation of the pattern. In this stimulus situation the optomotor responses for yaw, pitch and roll torque have about the same amplitudes and dynamic properties (Fig. 2). Pronounced negative responses are measured with periodic gratings of low pattern wavelengths due to geometrical interference (Fig. 3). The responses depend upon the contrast frequency rather than the angular velocity of the pattern (Fig. 4). Like the optomotor yaw response, roll and pitch responses can be elicited by small field motion in most parts of the visual field; only for motion below and behind the fly roll and pitch responses have low sensitivity.The mutantoptomotor-blind ^H31 (omb ^H31) in which the giant neurones of the lobula plate are missing or severely reduced, is impaired in all 3 optomotor torque responses (Fig. 5) whereas other visual responses like the optomotor lift/thrust response and the landing response (elicited by horizontal front-to-back motion) are not affected (Heisenberg et al. 1978).We propose that the lobula plate giant neurons mediate optomotor torque responses and that the VS-cells in particular are involved in roll and pitch but not in lift/thrust control. This hypothesis accommodates various electrophysiological and anatomical observations about these neurons in large flies.Abbreviation EMD elementary movement detector     

Adjustment of flight speed of gregarious desert locusts (Orthoptera: Acrididae) flying side by side

In tethered flying locusts, optomotor thrust responses induced by translatory pattern motion within the lateral visual fields were studied under closed-loop conditions. By modulating thrust in a compensatory manner, locusts counteracted a bias motion superposed on the thrust-related motion. This way, pattern speed was kept at 0° s^–1, indicating the set point of the respective optomotor control circuit. Though the quality of bias compensation varied greatly, it was largely independent from pattern characteristics. It might indicate that the gain of behavior not only is controlled by an automatic mechanism but also is affected by spontaneous modulations. Compensation of bias motion was critically dependent on the relation between self- and bias-generated motion: Locusts did not take control over pattern motion if self- and bias-generated motion differed greatly. Instead, locusts adopted a constant, supposingly preferred, thrust value. Therefore, flight speed is assumed to be controlled by two systems: the optomotor and a preferred thrust system. In free flight, an equalization of the flight speed of locusts within a swarm might result from similar behavior. In combination with a presumed coordination of the locusts' course direction, this may explain the continued cohesion of swarms in the field.     

Shared Visual Attention and Memory Systems in the Drosophila Brain

Background

Selective attention and memory seem to be related in human experience. This appears to be the case as well in simple model organisms such as the fly Drosophila melanogaster. Mutations affecting olfactory and visual memory formation in Drosophila, such as in dunce and rutabaga, also affect short-term visual processes relevant to selective attention. In particular, increased optomotor responsiveness appears to be predictive of visual attention defects in these mutants.

Methodology/Principal Findings

To further explore the possible overlap between memory and visual attention systems in the fly brain, we screened a panel of 36 olfactory long term memory (LTM) mutants for visual attention-like defects using an optomotor maze paradigm. Three of these mutants yielded high dunce-like optomotor responsiveness. We characterized these three strains by examining their visual distraction in the maze, their visual learning capabilities, and their brain activity responses to visual novelty. We found that one of these mutants, D0067, was almost completely identical to dunce¹ for all measures, while another, D0264, was more like wild type. Exploiting the fact that the LTM mutants are also Gal4 enhancer traps, we explored the sufficiency for the cells subserved by these elements to rescue dunce attention defects and found overlap at the level of the mushroom bodies. Finally, we demonstrate that control of synaptic function in these Gal4 expressing cells specifically modulates a 20–30 Hz local field potential associated with attention-like effects in the fly brain.

Conclusions/Significance

Our study uncovers genetic and neuroanatomical systems in the fly brain affecting both visual attention and odor memory phenotypes. A common component to these systems appears to be the mushroom bodies, brain structures which have been traditionally associated with odor learning but which we propose might be also involved in generating oscillatory brain activity required for attention-like processes in the fly brain.     

Developmental aspects of the innervation of skeletal muscle fibers in the chick embryo

Summary The M. complexus in the chick, commonly called the hatching muscle, undergoes conspicuous growth during the latter stages of embryonic development. Myogenesis of this muscle was compared to that of M. biceps femoris with regard to development of types of muscle fiber and their innervation. In both muscles fibers are of relatively uniform size and show little growth in diameter between 12 days of development and hatching; fibers develop continuously and display a wide range of diameters at all stages.Initial thickenings on the sarcolemma of fibers where axons are closely approximate were first observed at 10 days of development in both muscles. In both muscles fibers are innervated prior to fibers. Terminal axon networks bridge intercellular spaces and contact fibers in different myogenic clusters, fibers that develop on the surface membrane of fibers exhibit focal thickenings of the membrane and some cell projections that are directed toward axon- fiber contacts. These changes occurred only in fibers of M. complexus.At 14 days of embryogenesis, the processes of synaptogenesis and of myelin formation are less advanced in M. biceps femoris than in M. complexus. At this stage a fibers were observed to be innervated in M. complexus, but not yet in M. biceps femoris. Each fiber was observed to be encircled by several preterminal axons.It is concluded that the earlier development of M. complexus is correlated with an equally early development of nerve-muscle interactions.This work was supported in part by a grant from the Muscular Dystrophy Association of America, Inc.Postdoctoral Fellow of the Muscular Dystrophy Association I would like to thank Professor Dr. H. Tamate for his valuable advise. I am also grateful to Dr. L. Doerr, H. Stokes and Judi K. Lund for their advice and skilled technical assistance     

Are the large monopolar cells of the insect lamina on the optomotor pathway?

Evidence is presented here from experiments on the visual system of the fly that questions participation of the large monopolar cells (LMCs) in the optomotor response.

Reafferent control of optomotor yaw torque inDrosophila melanogaster

Summary In the flight simulator the optomotor response ofDrosophila melanogaster does not operate as a simple feedback loop. Reafferent and exafferent motion stimuli are processed differently. Under open-loop conditions responses to motion are weaker than under closed-loop conditions. It takes the fly less than 100 ms to distinguish reafferent from exafferent motion. In closed-loop conditions, flies constantly generate torque fluctuations leading to small-angle oscillations of the panorama. This reafferent motion stimulus facilitates the response to exafferent motion but does not itself elicit optomotor responses. Reafference control appears to be directionally selective: while a displacement of the patternm by as little as 0.1° against the expected direction leads to a fast syndirectional torque response, displacements in the expected direction have no comparable effect. Based on the behavior of the mutantrol sol, which under open-loop conditions is directionally motion-blind but in closed-loop conditions still performs optomotor balance, a model is proposed in which the fly's endogenous torque fluctuations are an essential part of the course control process. It is argued that the model may also account for wild type optomotor balance in the flight simulator.     

Recurrent inversion of visual orientation in the walking fly,Drosophila melanogaster

Summary Movement-induced visual orientation in flies depends largely upon predictable responses which establish simple optomotor balance or complex pseudo search in the appropriate visual environment. Less conspicuous course diverting spontaneous actions of the flies become important in pattern-induced visual orientation. The apparently stochastic spontaneous actions of the houseflyMusca domestica still allow powerful probabilistic predictions of orientation during stationary flight (Reichardt and Poggio 1981). The predominance of non-stochastic spontaneous actions such as body saccades, focussing and shift of visual attention, plasticity of response components etc. in the fruitflyDrosophila melanogaster (Heisenberg and Wolf 1979–1980) accounts for complementary behavioural options which reduce the relevance of probabilistic predictions of orientation in this fly.The conjecture of complementary options is based on a striking antagonism between orientation towards a visual object (fixation), and orientation in the opposite direction (anti-fixation), in the walking fly. Forced choice in a multiple-Y-maze quite definitely elicits fixation in the wild type, and antifixation in the optomotor blind mutantomb ^H31 (Fig. 3). However, these effects cannot be attributed to a continuous predominance of attraction in the wild type and repellence in the mutant. This is shown under comparable conditions of free choice in an arena: The flies of either strain alternate between fixation and anti-fixation of an inaccessible visual object (Fig. 4a), and keep running to and fro between two of these objects in Buridan's paradigm (Fig. 4b, c), even if the objects are not alike (Fig. 4d). The sequence of approach, retreat and transition may be repeated a few thousand times to the point of exhaustion (Fig. 5). The process resembles the recurrent alternation of ambiguous figures such as the Necker cube in human perception. The recurrent transition between competitive objects counteracts the accumulation of spontaneous preferences, and is likely to explain the apparent lack of pattern-discrimination under operant and non-operant conditions of continued free choice inDrosophila. The conspicuous dichotomy of fixation and anti-fixation in the same environment is, as yet, incompatible with the phenomenological theory of visually controlled orientation in larger flies.Abbreviation S.E.M. standard error of the mean     

Processing of figure and background motion in the visual system of the fly

The visual system of the fly is able to extract different types of global retinal motion patterns as may be induced on the eyes during different flight maneuvers and to use this information to control visual orientation. The mechanisms underlying these tasks were analyzed by a combination of quantitative behavioral experiments on tethered flying flies (Musca domestica) and model simulations using different conditions of oscillatory large-field motion and relative motion of different segments of the stimulus pattern. Only torque responses about the vertical axis of the animal were determined. The stimulus patterns consisted of random dot textures (Julesz patterns) which could be moved either horizontally or vertically. Horizontal rotatory large-field motion leads to compensatory optomotor turning responses, which under natural conditions would tend to stabilize the retinal image. The response amplitude depends on the oscillation frequency: It is much larger at low oscillation frequencies than at high ones. When an object and its background move relative to each other, the object may, in principle, be discriminated and then induce turning responses of the fly towards the object. However, whether the object is distinguished by the fly depends not only on the phase relationship between object and background motion but also on the oscillation frequency. At all phase relations tested, the object is detected only at high oscillation frequencies. For the patterns used here, the turning responses are only affected by motion along the horizontal axis of the eye. No influences caused by vertical motion could be detected. The experimental data can be explained best by assuming two parallel control systems with different temporal and spatial integration properties: TheLF-system which is most sensitive to coherent rotatory large-field motion and mediates compensatory optomotor responses mainly at low oscillation frequencies. In contrast, theSF-system is tuned to small-field and relative motion and thus specialized to discriminate a moving object from its background; it mediates turning responses towards objects mainly at high oscillation frequencies. The principal organization of the neural networks underlying these control systems could be derived from the characteristic features of the responses to the different stimulus conditions. The input to the model circuits responsible for the characteristic sensitivity of the SF-system to small-field and relative motion is provided by retinotopic arrays of local movement detectors. The movement detectors are integrated by a large-field element, the output cell of the network. The synapses between the detectors and the output cells have nonlinear transmission characteristics. Another type of large-field elements (pool cells) which respond to motion in front of both eyes and have characteristic direction selectivities are assumed to interact with the local movement detector channels by inhibitory synapses of the shunting type, before the movement detectors are integrated by the output cells. The properties of the LF-system can be accounted for by similar model circuits which, however, differ with respect to the transmission characteristic of the synapses between the movement detectors and the output cell; moreover, their pool cells are only monocular. This type of network, however, is not necessary to account for the functional properties of the LF-system. Instead, intrinsic properties of single neurons may be sufficient. Computer simulations of the postulated mechanisms of the SF-and LF-system reveal that these can account for the specific features of the behavioral responses under quite different conditions of coherent large-field motion and relative motion of different pattern segments.     

Expression of the C3-C4 intermediate character in somatic hybrids between Brassica napus and the C3-C4 species Moricandia arvensis

The wild crucifer Moricandia arvensis is a potential source of alien genes for the genetic improvement of related Brassica crops. In particular M. arvensis has a C₃-C₄ intermediate photosynthetic mechanism which results in enhanced recapture of photorespired CO₂ and may increase plant water-use efficiency. In order to transfer this trait into Brassica napus, somatic hybridisations were made between leaf mesophyll protoplasts from cultured M. arvensis shoot tips and hypocotyl protoplasts from three Brassica napus cultivars, Ariana, Cobra and Westar. A total of 23 plants were recovered from fusion experiments and established in the greenhouse. A wide range of chromosome numbers were observed among the regenerated plants, including some apparent mixoploids. Thirteen of the regenerated plants were identified as nuclear hybrids between B. napus and M. arvensis on the basis of isozyme analysis. The phenotypes of these hybrids were typically rather B. napus-like, but much variability was observed, including variation in flower colour, leaf shape and colour, leaf waxiness, fertility and plant vigour. CO₂ compensation point measurements on the regenerated plants demonstrated that 3 of the hybrids express the M. arvensis C₃-C₄ intermediate character at the physiological level. Semi-thin sections through leaf tissues of these 3 plants revealed the presence of a Kranz-like leaf anatomy characteristic of M. arvensis but not found in B. napus. This is the first report of the expression of this potentially important agronomic trait, transferred from Moricandia, in M. arvensis x B. napus hybrids.     

Visual control of straight flight in Drosophila melanogaster

Summary The optomotor system of Drosophila is investigated in a flight simulator in which the fly's yaw torque controls the angular velocity of the panorama (striped drum, negative feedback). Flies in the flight simulator maintain a stable orientation even in a homogeneously textured panorama without landmarks. During straight flight, torque is not zero. It consists of small pulses mostly alternating in polarity. The course is controlled by the duration (and possibly amplitude) of the pulses. The system operates under reafference control. By comparing the pulses with the visual input the system continuously measures and adjusts the efficacy of the torque output. The comparison, however, is not between angular velocity and yaw torque but, instead, between visual acceleration and pretorque, the first time derivative of torque. For comparison, the system first computes a cross-correlation. If the correlation coefficient is above a certain threshold the system calculates the external gain and adjusts its internal gain so as to keep the total gain constant. With the correlation coefficient below threshold, however, the system keeps the internal gain low despite the infinitely small external gain. We propose that for a reafferent optomotor system the coupling coefficient and the correlation coefficient of pretorque and visual acceleration are more relevant than the distinction between exafference and reafference.Abbreviation EMD elementary movement detector     

Kinetic properties of bovine brain proteinl-isoaspartyl methyltransferase determined using a synthetic isoaspartyl peptide substrate

Proteinl-isoaspartyl methyltransferase, an enzyme enriched in brain, is implicated in the repair of age-damaged proteins containing atypical, isoaspartyl peptide bonds. We have investigated the kinetics of methylation using a synthetic peptide substrate having the structure Trp-Ala-Gly-Gly-isoAsp-Ala-Ser-Gly-Glu. Double-reciprocal plots of initial velocity versus concentration of S-adenosylmethionine (AdoMet) at different fixed concentrations of peptide gave straight lines converging at a positive 1/v value and a negative 1/AdoMet value. The product S-adenosylhomocysteine (AdoHcy) was a competitive inhibitor towards AdoMet and a linear mixed-type inhibitor towards peptide. These results are consistent with the rapid-equilibrium random sequential bi-bi mechanism previously proposed for the enzyme, but they also reveal the formation of the deadend, enzyme-peptide-AdoHcy, complex. The rate constants were:V _max=32–34 nmol/min/mg,K ^peptide=7.6–9.4 M,K ^AdoMet=1.9–2.2 M, =0.43–0.53,K ^AdoHcy=0.08 M, =2.9. The interaction factors and indicate that binding of enzyme to peptide increases its affinity for AdoMet and decreases its affinity for AdoHcy. Methylation was linear with time throughout the transfer of 2 mol of methyl groups/mol of enzyme. This absence of burst kinetics suggests that slow release of products cannot explain the low turnover number.Special issue dedicated to Dr. Paul Greengard     

Motion parallax and figural properties of depth control flight speed in an insect

Male gypsy moths (Lymantria dispar L.) are able to control their forward flight speed solely by means of optical cues derived from the lateral parts of their visual field. Thereby, relative motion of the objects is required, which under free flight conditions would derive from the self-induced motion of the stationary objects within a surrounding structured in depth. Besides this motion parallax, however, the control system for forward flight speed demands figural properties of the objects such as differences in their angular extension or contrast. In a natural surround, the images of objects closer to the moth have higher angular velocity and, in addition, are usually larger and have higher contrast than those of objects farther away. The experiments have shown that this natural pairing of angular velocity and figural properties is essential to induce a thrust response which appears suitable to counteract involuntary changes in forward flight speed, as e.g. caused by changes in wind speed under free flight conditions. Inverse pairing of the visual stimuli caused the moths to either enlarge the error signal according to positive feedback within the control circuit, or to increase thrust to a maximum as always found in experiments without motion parallax or in the absence of differences in figural properties. The open-loop experiments furthermore established that the set point of angular velocity of the closer objects lies within the range of 4.5–9°/s, and that pattern speed modulation has to cover this range in order to induce a compensatory thrust modulation. The response is largely independent of the magnitude of relative motion as long as it is present; the response amplitude, however, strongly depends on the amplitude of pattern speed modulation.The results are discussed with regard to other visual cues used for the control of forward flight speed in insects, the algorithm underlying the figure-ground discrimination in flies, and the perception of depth in man as known from psychophysical experiments.     

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).     

Cell calcium signalling induced by endogenous lectin carbohydrate interaction in the Jurkat T cell line

The effects of the -galactoside-binding lectin from human placenta (HPL14) on intracellular calcium concentration ([Ca²⁺]_i) were examined in the human Jurkat T cell line. The lectin induces a concentration dependent increase in [Ca²⁺]_i. This calcium signalling effect is clearly mediated through complementary cell surface galactoglycoconjugates because it can be blocked by -galactosides. The observed Ca²⁺-response involves both the release of calcium from intracellular stores and a calcium influx from the extracellular space. It is sustained in the presence of 1 mM extracellular calcium whereas it becomes transient when the influx of extracellular calcium was blocked by calcium chelation to EGTA. Voltage-sensitive calcium channel blockers like verapamil and prenylamine were without effect on the action of HPL14. Protection of the sugar binding activity of HPL14 in the absence of a thiol-reducing reagent by carboxamidomethylation (CM-HPL14) or by substitution Cys2 with serine (C2S) results in lectin proteins with considerably decreased calcium signalling efficiency. The recombinant lectin (Rec H) and the mutant protein obtained by substitution of highly conservative Trp68 with tyrosine (W68Y) induce lower levels of [Ca²⁺]_i compared to wild type lectin.Abbreviation [Ca²⁺]_i concentration of intracytoplasmic free calcium - CM carboxamidomethylation - CRD earbohydrate recognition domain - C2S mutant lectin protein in which Cys2 was replaced by serine - EGTA ethyleneglycol-bis(2-aminoethylether)-N,N,N - N-tetraacetic acid HEPES,N-(2-hydroxyethyl)piperazine-N-2-ethanesulfonic acid - HPL14 human -galactoside-binding placental lectin - Rec H recombinant human 14 kDa lectin - W68Y mutant lectin protein in which Trp68 was substituted to tyrosine     

Die Hemmung der Phytochrominduzierten Photomorphogenese („Positive” Photomorphosen) des Senfkeimlings (Sinapis alba L.) Durch Actinomycin D und Puromycin

Zusammenfassung Die positiven Photomorphosen Öffnung des Hypokotylhakens und Entfaltung der Kotyledonen können ganz ähnlich wie die phytochrominduzierte Anthocyansynthese und andere positive Photomorphosen durch Actinomycin D und Puromycin gehemmt werden. Man kann daraus schließen, daß diese beiden photomorphogenetischen Reaktionen des Senfkeimlings ebenfalls durch eine von P₇₃₀ über eine Signalkette ausgelöste Aktivierung von potentiell aktiven Genen veranlaßt werden.

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The inhibition of phytochrome-mediated photomorphogenesis (positive photoresponses) by actinomycin D and puromycin in the mustard seedling (Sinapis alba L.)

Summary The many photochrome-mediated photoresponses of a seedling (Sinapis alba L., white seeded mustard) can be divided into 3 categories: positive, negative, and complex photoresponses. Positive photoresponses are those which are characterized by an initiation or a promotion of biosynthetic or growth processes (Mohr, 1966b). Phytochrome-mediated anthocyanin synthesis is the prototype of a positive photoresponse. It has been shown in previous papers (e.g. Lange and Mohr, 1965; Mohr et al., 1965) that positive photoresponses can be specifically inhibited by actinomycin D and puromycin. It has been concluded that in the case of positive photoresponses P₇₃₀ (the active phytochrome) exerts its function through differential gene activation.—In the present paper it has been demonstrated that phytochrome-mediated positive photoresponses of the mustard seedling like opening of the hypocotylar hook and unfolding of the cotyledons can be inhibited by relatively low doses of actinomycin D and puromycin in very much the same way as anthocyanin synthesis or cotyledon enlargement is inhibited. It has been concluded that in these cases too the action of P₇₃₀ must be attributed to an activation of potentially active genes in the manner postulated on the basis of the data on anthocyanin synthesis.

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Die Arbeit wurde durch Sachbeihilfen der Deutschen Forschungsgemeinschaft und der Stiftung Volkswagenwerk (an Prof. Mohr) ermöglicht.     

Comparison between the movement detection systems underlying the optomotor and the landing response in the housefly

Flies evaluate movement within their visual field in order to control the course of flight and to elicit landing manoeuvres. Although the motor output of the two types of responses is quite different, both systems can be compared with respect to the underlying movement detection systems. For a quantitative comparison, both responses were measured during tethered flight under identical conditions. The stimulus was a sinusoidal periodic pattern of vertical stripes presented bilaterally in the fronto-lateral eye region of the fly. To release the landing response, the pattern was moved on either side from front to back. The latency of the response depends on the stimulus conditions and was measured by means of an infrared light-beam that was interrupted whenever the fly lifted its forelegs to assume a preprogrammed landing posture (Borst and Bahde 1986). As an optomotor stimulus the pattern moved on one side from front to back and on the other side in the opposite direction. The induced turning tendency was measured by a torque meter (Götz 1964). The response values which will be compared are the inverse latencies of the landing response and the amplitude of the yaw torque.

1. Optomotor course-control is more sensitive to pattern movement at small spatial wavelengths (10° and 20°) than the landing response (Fig. 1a and b). This suggests that elementary movement detectors (EMDs, Buchner 1976) with large detection base (the distance between interacting visual elements) contribute more strongly to the landing than to the optomotor system.
2. The optimum contrast frequencies of the different responses obtained at a comparatively high pattern contrast of about 0.6 was found to be between 1 and 10 Hz for the optomotor response, and around 20 Hz for the landing response (Fig. 2a and b). This discrepancy can be explained by the fact that the optomotor response was tested under stationary conditions (several seconds of stimulation) while for the landing response transient response characteristics of the movement detectors have to be taken into account (landing occurs under these conditions within less than 100 ms after onset of the movement stimulus). To test the landing system under more stationary conditions, the pattern contrast had to be reduced to low values. This led to latencies of several seconds. Then the optimum of the landing response is around 4 Hz. This is in the optimum range of the optomotor course-control response. The result suggests the same filter time constants for the movement detectors of both systems.
3. The dependence of both responses on the position and the size of the pattern was examined. The landing response has its optimum sensitivity more ventrally than the optomotor response (Fig. 3a and b). Both response amplitudes increase with the size of the pattern in a similar progression (Fig. 3c and d).

In first approximation, the present results are compatible with the assumption of a common set of movement detectors for both the optomotor course-control and the landing system. Movement detectors with different sampling bases and at different positions in the visual field seem to contribute with different gain to both responses. Accordingly, the control systems underlying both behaviors are likely to be independent already at the level of spatial integration of the detector output.     

First Field Studies of an Endangered South African seahorse, Hippocampus capensis

South Africa's endemic Knysna seahorse, Hippocampus capensis Boulenger 1900, is a rare example of a marine fish listed as Endangered by the IUCN because of its limited range and habitat vulnerability. It is restricted to four estuaries on the southern coast of South Africa. This study reports on its biology in the Knysna and Swartvlei estuaries, both of which are experiencing heavy coastal development. We found that H. capensis was distributed heterogeneously throughout the Knysna Estuary, with a mean density of 0.0089m^–2 and an estimated total population of 89000 seahorses (95% confidence interval: 30000–148000). H. capensis was found most frequently in low density vegetation stands (20% cover) and grasping Zostera capensis. Seahorse density was not otherwise correlated with habitat type or depth. The size of the area in which any particular seahorse was resighted did not differ between males and females. Adult sex ratios were skewed in most transects, with more males than females, but were even on a 10 by 10m focal study grid. Only three juveniles were sighted during the study. Both sexes were reproductively active but no greeting or courtship behaviours were observed. Males on the focal study grid were longer than females, and had shorter heads and longer tails, but were similar in colouration and skin filamentation. The level of threat to H. capensis and our limited knowledge of its biology mean that further scientific study is urgently needed to assist in developing sound management practices.     

What kind of movement detector is triggering the landing response of the housefly?

As shown before, the latency of the housefly's landing response depends on the conditions of the visual stimulus (Borst 1986). Accordingly, the latency can be used to characterize the movement detection system which is triggering the landing response.The stimulus was a sinusoidal periodic pattern of vertical stripes presented bilaterally in the frontolateral eye region of the fly. It started to move, simultaneously on either side, from front to back at a given time. The latency of the response was measured by means of an infrared light-beam that was interrupted whenever the fly lifted its forelegs to assume a preprogrammed landing posture (Fig. 1). The latency was found to vary in a range from 60 ms up to several seconds depending on the pattern's spatial wavelength , contrast frequency cf and contrast C.For sufficiently high pattern contrast the optimum of the reaction (minimum latency) is found at spatial wavelengths of 30–40° and contrast frequencies of 8–17 periods/s (Fig. 3a). This is about 2–10 times more than is anticipated from the optomotor response under similar conditions. Evaluation of the optimum contrast frequency cf _OPT at different wavelengths shows that cf _OPT is not independent of (Fig. 3b, solid line). The same is true for the contrast dependence of the reaction: reduction of the contrast leads not only to a general decrease in the response amplitudes (prolongation of the latency) (Fig. 4a), but also to a shift of cf _OPT towards lower contrast frequencies (Fig. 4b, solid line).In the theory of the correlation-type movement detector (Reichardt 1961) which underlies the optomotor response of flies the dependence of cf _OPT on pattern wavelength and/or pattern contrast is not expected under stationary conditions. However, as shown by computer simulation all experimental results can be explained by a homogeneous retinotopic array of correlation movement detectors (Fig. 2) if their response under non-stationary conditions is taken into account. We simply assume that the spatially and temporally integrated output of the movement detectors is evaluated by a threshold device (Fig.5). The correlation-type movement detection in combination with a temporal integrator system predicts the rather complex dependence of the optimum contrast frequency on pattern wavelength and pattern contrast (dashed lines in Fig. 3b and 4b) and provides the missing explanation of the variable latencies of the landing response.Comparing the parameters of the correlation-type movement detector derived in the present study with those of the optomotor response, the landing response seems to use the same type of movement detection system. To account for the high wavelength optimum, however, the input elements of the movement detection system of the landing response might have an increased visual field (e.g. by pooling neighbouring visual elements) and, accordingly, a reduced visual acuity as compared with the input elements of the optomotor system.Abbreviations (°) spatial pattern wavelength - w(°/s) angular velocity of the pattern - cf (Hz) contrast frequency=w/ - cf _OPT(Hz) cf leading to the shortest latency - (Hz) angular frequency=2cf - I mean luminance of the pattern - I modulation amplitude of the pattern - C pattern contrast=I/ - (ms) time constant of a filter - (°) angle between the optical axis of neighbouring visual elements - (°) acceptance angle of visual elements     

Cholinesterases and cell proliferation in “nonstratified” and “stratified” cell aggregates from chicken retina and tectum

Summary Dissociated single cells from chicken retina or tectum kept in rotation-mediated cell culture aggregate, proliferate and establish a certain degree of histotypical cellto-cell relationships (sorting out), but these systems never form highly laminated aggregates (nonstratified R- and T-aggregates). In contrast, a mixture of retinal plus pigment epithelial cells forms highly stratified aggregates (RPE-aggregates, see Vollmer et al. 1984). The present comparative study of stratified and nonstratified aggregates enables us to investigate the process of cell proliferation uncoupled from that of tissue stratification. Here we try to relate these two basic neurogenetic processes with patterns of expression of cholinesterases (AChE, BChE) during formation of both types of aggregates.During early aggregate formation, in both stratified and nonstratified aggregates an increased butyrylcholinesterase activity is observed close to mitotically active cells. Quantitatively both phenomena show their maxima after 2–3 days in culture. In contrast, AChE-expression in all systems increases with incubation time. In nonproliferative areas, in the center of RPE-aggregates, the formation of plexiform layers is characterized initially by weak BChE and then strong AChE-activity. These areas correspond with the inner (IPL) and outer (OPL) plexiform layers of the retina in vivo. Although by sucrose gradient centrifugation we find that the 6S- and the fiber-associated 11S-molecules of AChE are present in all types of aggregates, during the culture period the ratio of 11S/6S-forms increases only in RPE-aggregates, which again indicates the advanced degree of differentiation within these aggregates.It is thus demonstrated that cholinesterases first correlate with neuronal cell proliferation and later with stratification, which indicates functions of both enzymes during both developmental periods.Abbreviations AChE acetylcholinesterase - BChE butyrylcholinesterase - iso-OMPA specific inhibitor of BChE - BW 284C51 specific inhibitor of AChE - IPL inner plexiform layer - OPL outer plexiform layer     

Influence of Nematode Parasitism,Body Size,Temperature, and Diel Period on the Flight Capacity of <Emphasis Type="Italic">Sirex noctilio</Emphasis> F. (Hymenoptera: Siricidae)

Sirex noctilio F. (Hymenoptera: Siricidae) is a woodwasp of pine trees that has recently invaded and established in North American forests. Although S. noctilio has had a limited impact in North America to date, there is some concern that it could have a significant impact on pine plantations, especially in the southeastern U.S.A. Moreover, there are few data on the flight capacity of male S. noctilio. We found no association between parasitism by D. siricidicola and whether or not S. noctilio initiated flight on the flight mill. Male wasps that were parasitized by nematodes were heavier than non-parasitized males, but there was no significant difference in mass between parasitized and non-parasitized females. We also examined the flight capacity of male and female S. noctilio in relation to nematode parasitism, body mass, temperature (for only males), and diel period. Body mass, temperature, and diel period affected flight in S. noctilio such that wasps were generally observed to fly faster, farther, and more frequently if they were heavier, flying at warmer temperatures, and flying during the photoperiod. The fact that nematode-parasitized male wasps were found to fly farther than the non-parasitized males is consistent with the hypothesis that nematode parasitism does not negatively affect the flight capacity of S. noctilio.