Phototaxis and photophobic responses in Stentor coeruleus action spectrum and role of Ca2+ fluxes

Negative phototactic orientation, step-up photophobic responses and light-induced action potentials have been studied in the ciliate Stentor coeruleus. A resolved action spectrum, based on fluence rate-response curves, is consistent with stentorin as the photoreceptor. Calcium flux blockers prolong the response time for ciliary stop and reversal and inhibit step-up photophobic responses. Drugs believed to affect the membrane-bound calcium pump likewise inhibit phobic responses. On the other hand, α-phosphatidic acid promotes Ca²⁺-influx and enhances the photophobic sensitivity of the organism, thus providing an unambiguous evidence for the role of Ca²⁺ influx. A change in the response time decreases the degree of phototactic orientation, indicating that negative phototaxis in this organism is brought about by subsequent phobic responses of individual rows of cilia as the associated photoreceptor granules experience an increase in light intensity when the organism rotates during forward locomotion in lateral light.     

Formation of the function of the photosensing system in early development

The function of simple eyes in two planarian species, two-eyed Girardia tigrina and multi-eyed Polycelis tenuis, has been studied. When exposed to light, planarians display a light avoidance reaction known as negative phototaxis. This reaction has been investigated in intact animals and in head and tail fragments in the course of eye regeneration after their section. Specific features of the phototaxis reaction have been described in all groups of animals. The differences in light response recovery were shown between two planarian species and two regenerating fragments. No correlation has been found between phototactic reactions and restoration of eye structure, the number of eyes, maturation of the ganglion, growth of regenerative blastema, and motor system. The phototactic response occurred two days after the recovery of the morphology of eyes and their connection with the brain. The participation of conserved and novel genes in early development of the eye function is discussed.     

Effect of external ionic environment on phototaxis of Volvox carteri

The sign of phototaxis in Volvox carteri is temperature-dependent;positive at room temperature and negative at low temperature.Modification of the tactic sign by external ions, pH and chemicalswas studied. The addition of 30 mM potassium ion to the mediumchanged the tactic sign from positive to negative, and a mediumwith a high pH elicited positive phototaxis. An increase inthe potassium or hydrogen ion concentration raised the reversaltemperature of the phototactic sign, and the addition of magnesiumor calcium ion also raised the reversal temperature of the signslightly. Valinomycin, a highly specific ionophore of potassium,raised the reversal temperature. CCCP, DCMU and DCCD, whichdepolarize biological membranes, also raised the reversal temperature.These results show that the sign of phototaxis is determinedby membrane polarization; on depolarization of the membranethe sign of phototaxis changes from positive to negative. Ethanol lowered the reversal temperature, and sodium azide inhibitedboth positive and negative phototaxis. The effects of ethanoland azide indicate that depolarization of the membrane is notthe only factor that induces change in the phototactic sign. (Received July 23, 1979; )     

Evolution of phototaxis

Phototaxis in the broadest sense means positive or negative displacement along a light gradient or vector. Prokaryotes most often use a biased random walk strategy, employing type I sensory rhodopsin photoreceptors and two-component signalling to regulate flagellar reversal. This strategy only allows phototaxis along steep light gradients, as found in microbial mats or sediments. Some filamentous cyanobacteria evolved the ability to steer towards a light vector. Even these cyanobacteria, however, can only navigate in two dimensions, gliding on a surface. In contrast, eukaryotes evolved the capacity to follow a light vector in three dimensions in open water. This strategy requires a polarized organism with a stable form, helical swimming with cilia and a shading or focusing body adjacent to a light sensor to allow for discrimination of light direction. Such arrangement and the ability of three-dimensional phototactic navigation evolved at least eight times independently in eukaryotes. The origin of three-dimensional phototaxis often followed a transition from a benthic to a pelagic lifestyle and the acquisition of chloroplasts either via primary or secondary endosymbiosis. Based on our understanding of the mechanism of phototaxis in single-celled eukaryotes and animal larvae, it is possible to define a series of elementary evolutionary steps, each of potential selective advantage, which can lead to pelagic phototactic navigation. We can conclude that it is relatively easy to evolve phototaxis once cell polarity, ciliary swimming and a stable cell shape are present.     

Multiple light inputs control phototaxis in Synechocystis sp. strain PCC6803

The phototactic behavior of individual cells of the cyanobacterium Synechocystis sp. strain PCC6803 was studied with a glass slide-based phototaxis assay. Data from fluence rate-response curves and action spectra suggested that there were at least two light input pathways regulating phototaxis. We observed that positive phototaxis in wild-type cells was a low fluence response, with peak spectral sensitivity at 645 and 704 nm. This red-light-induced phototaxis was inhibited or photoreversible by infrared light (760 nm). Previous work demonstrated that a taxD1 mutant (Cyanobase accession no. sll0041; also called pisJ1) lacked positive but maintained negative phototaxis. Therefore, the TaxD1 protein, which has domains that are similar to sequences found in both bacteriophytochrome and the methyl-accepting chemoreceptor protein, is likely to be the photoreceptor that mediates positive phototaxis. Wild-type cells exhibited negative phototaxis under high-intensity broad-spectrum light. This phenomenon is predominantly blue light responsive, with a maximum sensitivity at approximately 470 nm. A weakly negative phototactic response was also observed in the spectral region between 600 and 700 nm. A deltataxD1 mutant, which exhibits negative phototaxis even under low-fluence light, has a similar action maximum in the blue region of the spectrum, with minor peaks from green to infrared (500 to 740 nm). These results suggest that while positive phototaxis is controlled by the red light photoreceptor TaxD1, negative phototaxis in Synechocystis sp. strain PCC6803 is mediated by one or more (as yet) unidentified blue light photoreceptors.     

Phototaxis by the Dinoflagellate Gymnodinium splendens Lebour*

A microscope-television system was used to monitor quantitatively the behavior of Gymnodinium splendens Lebour in response to light. The predominant behavioral sequence upon stimulation is (a) an initial 2–5 sec cessation of movement (stop-response) followed by (b) positive phototaxis. The action spectra for each response are identical, having maxima at 450 and 280 nm. Upon measuring the percent response to a range of stimulus intensities, it is apparent that a stop-response is not a behavioral prerequisite for phototaxis. An identical circadian rhythm in photoresponsiveness is observed for phototaxis and for the stop-response with greatest light sensitivity occurring during the first 4 hr of the entrained light period. The implication of phototactic sensitivity and the phototactic circadian rhythm in diurnal vertical migration is discussed.     

How 5000 independent rowers coordinate their strokes in order to row into the sunlight: Phototaxis in the multicellular green alga <Emphasis Type="Italic">Volvox</Emphasis>

Background

The evolution of multicellular motile organisms from unicellular ancestors required the utilization of previously evolved tactic behavior in a multicellular context. Volvocine green algae are uniquely suited for studying tactic responses during the transition to multicellularity because they range in complexity from unicellular to multicellular genera. Phototactic responses are essential for these flagellates because they need to orientate themselves to receive sufficient light for photosynthesis, but how does a multicellular organism accomplish phototaxis without any known direct communication among cells? Several aspects of the photoresponse have previously been analyzed in volvocine algae, particularly in the unicellular alga Chlamydomonas.

Results

In this study, the phototactic behavior in the spheroidal, multicellular volvocine green alga Volvox rousseletii (Volvocales, Chlorophyta) was analyzed. In response to light stimuli, not only did the flagella waveform and beat frequency change, but the effective stroke was reversed. Moreover, there was a photoresponse gradient from the anterior to the posterior pole of the spheroid, and only cells of the anterior hemisphere showed an effective response. The latter caused a reverse of the fluid flow that was confined to the anterior hemisphere. The responsiveness to light is consistent with an anterior-to-posterior size gradient of eyespots. At the posterior pole, the eyespots are tiny or absent, making the corresponding cells appear to be blind. Pulsed light stimulation of an immobilized spheroid was used to simulate the light fluctuation experienced by a rotating spheroid during phototaxis. The results demonstrated that in free-swimming spheroids, only those cells of the anterior hemisphere that face toward the light source reverse the beating direction in the presence of illumination; this behavior results in phototactic turning. Moreover, positive phototaxis is facilitated by gravitational forces. Under our conditions, V. rousseletii spheroids showed no negative phototaxis.

Conclusions

On the basis of our results, we developed a mechanistic model that predicts the phototactic behavior in V. rousseletii. The model involves photoresponses, periodically changing light conditions, morphological polarity, rotation of the spheroid, two modes of flagellar beating, and the impact of gravity. Our results also indicate how recently evolved multicellular organisms adapted the phototactic capabilities of their unicellular ancestors to multicellular life.

     

Perching of Tengmalm's Owl (Aegolius funereus) Nestlings at the Nest Box Entrance: Effect of Time of the Day,Age, Wing Length and Body Weight

The behaviour of the nestlings of nocturnal cavity-nesting species has relatively rarely been studied in detail because of problems connected with use of the technical devices required to provide long-term monitoring of individuals. However, long-term observation of nestling behaviour is crucial in order to identify different types of behaviour which may be caused by sibling competition at the end of nesting period. We studied behaviour of 43 Tengmalm''s owl (Aegolius funereus) nestlings at 14 nests using a camera and a chip system. The nestlings perched at the nest box entrance from an average age of 28 days from hatching (range 24–34 days) until fledging, spending around 2 hours per day here in total, in periods ranging from a few seconds to 147 min (7.6±10.9 min, mean ± SD). We found that individual duration of perching at the nest box entrance was significantly influenced by nestlings'' age and wing length and that the duration of perching at the nest box entrance significantly decreased with time of night. However, during daylight hours, time of day had no effect on either probability or duration of nestlings'' perching. We suggest daylight perching at the nest box entrance results from nestlings'' preparation for fledging, while individuals perching here during the night may gain an advantageous position for obtaining food from the parents; another possibility at all times of day is that nestlings can reaffirm their social dominance status by monopolizing the nest box entrance.     

Phototaxis and membrane potential in the photosynthetic bacterium Rhodospirillum rubrum.

Cells of the photosynthetic bacterium Rhodospirillum rubrum cultivated anaerobically in light show phototaxis. The behavior of individual cells in response to the phenomenon is reversal(s) of the swimming direction when the intensity of the light available to them abruptly decreases. The tactic response was inhibited by antimycin, an inhibitor of the photosynthetic electron transfer system. The inhibitory effect of antimycin was overcome by phenazine methosulfate. Motility of the cells was not impaired by antimycin under aerobic conditions. Valinomycin plus potassium also inhibited their phototactic response; however, valinomycin or potassium alone had no effect. A change in membrane potential of the cells was measured as an absorbance change of carotenoid. Changes in the membrane potential caused by "on-off" light were prevented by antimycin and by valinomycin plus potassium, but not by antimycin plus phenazine methosulfate nor valinomycin or potassium alone. The results indicated that the phototactic response of R. rubrum is mediated by a sudden change in electron flow in the photosynthetic electron transfer system, and that the membrane potential plays an important role in manifestation of the response.     

Morphological and functional recovery of the planarian photosensing system during head regeneration

When exposed to light, planarians display a distinctive light avoidance behavior known as negative phototaxis. Such behavior is temporarily suppressed when animals are decapitated, and it is restored once the animals regenerate their heads. Head regeneration and the simple but reproducible phototactic response of planarians provides an opportunity to study the association between neuronal differentiation and the establishment of behavior in a simple, experimentally tractable metazoan. We have devised a phototaxis assay system to analyze light response recovery during head regeneration and determined that light evasion is markedly re-established 5 days after amputation. Immunohistological and in situ hybridization studies indicate that the photoreceptors and optic nerve connections to the brain begin by the fourth day of cephalic regeneration. To experimentally manipulate the light response recovery, we performed gene knockdown analysis using RNA interference (RNAi) on two genes (1020HH and eye53) previously reported to be expressed at 5 days after amputation and in the dorso-medial region of the brain (where the optic nerves project). Although RNAi failed to produce morphological defects in either the brain or the visual neurons, the recovery of the phototactic response normally observed in 5-day regenerates was significantly suppressed. The data suggest that 1020HH and eye53 may be involved in the functional recovery and maintenance of the visual system, and that the phototaxis assay presented here can be used to reliably quantify the negative phototactic behavior of planarians.     

Deuterium oxide (D2O) enhances the photosensitivity of Stentor coeruleus.

Stentor coeruleus exhibits negative phototaxis and step-up photophobic response (avoiding reaction) to visible light (maximum at 610-620 nm in both responses). In the presence of deuterium oxide (D2O) the step-up photophobic response was markedly enhanced, whereas the phototactic orientation response was inhibited. The induction time for the step-up photophobic response was longer in D2O than in H2O, and the duration of ciliary reversal for the response was also longer in D2O than in H2O, indicating that certain steps of the sensory transduction chain are subject to solvent deuterium isotope effects. The enhancement of the step-up photophobic response in D2O was canceled by LaCl3, while the inhibition of the phototactic orientation response in D2O was partially removed by LaCl3, even though LaCl3 did not affect the phototactic orientation response. These results suggest that the sensory transduction mechanisms for the two photoresponses are different, although the photoreceptors (stentorin) are the same.     

光谱和光强度对西花蓟马雌虫趋光行为的影响

利用行为学方法研究了光谱、光强对西花蓟马Frankliniella occidentalis (Pergande)雌成虫的趋、避光行为的影响。结果显示：（1）在340~605nm波谱内14个波长其光谱趋光行为反应为多峰型,峰间主次较明显。趋光行为反应中,蓝绿区498~524nm有一较宽峰,趋光率20.31%;其它各峰依大小次序分别位于紫光380nm、蓝光440nm;（２）避光行为反应中,蓝光440nm处略高,避光率17.19%;紫外340nm处亦有一峰,避光率15.63%;（３）随光强增强其趋光反应率增大,白光、380nm和524nm刺激时其光强趋光行为反应呈一倒“L”型式样,498nm为峰型,440nm时为一较缓的平直线型;光强最弱时仍均有一定趋光率 ,最强时均未出现高端平台;（４）随光强增强其避光反应率增大,440nm为较平缓直线;340nm刺激时为较缓波动线。结果表明：光谱对其趋光行为有很大影响,光强度的影响较大且影响大小与波长因素有关。     

The Process and Causes of Fledging in a Cavity-Nesting Passerine Bird, the House Wren (Troglodytes aedon)

Little is known about the process or causes of fledging or nest‐leaving in passerine birds because researchers can rarely predict when fledging will occur in a given nest. We used continuous videotaping of nests to both document the process of fledging in the house wren, Troglodytes aedon, a small, cavity‐nesting songbird, and test hypotheses as to what might cause fledging to begin. Fledging began any time from 14 to 19 d after hatching commenced. Slower‐developing broods fledged later than faster‐developing broods. Fledging typically began within 5 h of sunrise and over 80% of all nestlings fledged before noon. All nestlings fledged on the same day at 65% of nests and over two consecutive days in most other nests. We found no evidence that fledging was triggered by changes in parental behaviour. Parental rate of food delivery to nestlings did not decline during a 3‐h period leading up to the first fledging, nor was the rate of feeding just prior to the first fledging lower than the rate at the same time the day before. Moreover, parents did not slow the rate of food delivery to nests after part of the brood had fledged. Hatching is asynchronous in our study population which creates a marked age/size hierarchy within broods. At most nests, the first nestling to fledge was the most well‐developed nestling in the brood or nearly so (as measured by feather length). This suggests that fledging typically begins when the most well‐developed nestlings in the brood reach some threshold size. However, at about one‐fifth of nests, the first nestling to fledge was only moderate in size. At these nests, severe competition for food may have caused smaller, less competitive nestlings to fledge first to increase their access to food. We found no strong support for the suggestion that the oldest nestlings delay fledging until their least‐developed nestmate reaches some minimum size, although further experimental work on this question is warranted.     

The photobehaviour of Daphnia spp. as a model to explain diel vertical migration in zooplankton

Many pelagic animal species in the marine environment and in lakes migrate to deeper water layers before sunrise and return around sunset. The amplitude of these diel vertical migrations (DVM) varies from several hundreds of metres in the oceans to approx. 5–20 m in lakes. DVM can be studied from a proximate and an ultimate point of view. A proximate analysis is intended to reveal the underlying behavioural mechanism and the factors that cause the daily displacements. The ultimate analysis deals with the adaptive significance of DVM and the driving forces that were responsible for the selection of the traits essential to the behavioural mechanism. The freshwater cladoceran Daphnia is the best studied species and results can be used to model migration behaviour in general. Phototaxis in Daphnia spp., which is defined as a light-oriented swimming towards (positive phototaxis) or away (negative phototaxis) from a light source, is considered the most important mechanism basic to DVM. A distinction has been made between primary phototaxis which occurs when light intensity is constant, and secondary phototaxis which is caused by changes in light intensity. Both types of reaction are superimposed on normal swimming. This swimming of Daphnia spp. consists of alternating upwards and downwards displacements over small distances. An internal oscillator seems to be at the base of these alternations. Primary phototaxis is the result of a dominance of either the upwards or the downwards oscillator phase, and the direction depends on internal and external factors: for example, fish-mediated chemicals or kairomones induce a downwards drift. Adverse environmental factors may produce a persistent primary phototaxis. Rare clones of D. magna have been found that show also persistent positive or negative primary phototaxis and interbreeding of the two types produces intermediate progeny: thus a genetic component seems to be involved. Also secondary phototaxis is superimposed on normal swimming: a continuous increase in light intensity amplifies the downwards oscillator phase and decreases the upwards phase. A threshold must be succeeded which depends on the rate and the duration of the relative change in light intensity. The relation between both is given by the stimulus strength versus stimulus duration curve. An absolute threshold or rheobase exists, defined as the minimum rate of change causing a response if continued for an infinitely long time. DVM in a lake takes place during a period of 1-5-2 h when light changes are higher than the rheobase threshold. Accelerations in the rate of relative increase in light intensity strongly enhance downwards swimming in Daphnia spp. and this enhancement increases with increasing fish kairomone and food concentration. This phenomenon may represent a ‘decision-making mechanism’ to realize the adaptive goal of DVM: at high fish predator densities, thus high kairomone concentrations, and sufficiently high food concentrations, DVM is profitable but not so at low concentrations. Body axis orientation in Daphnia spp. is controlled with regard to light-dark boundaries or contrasts. Under water, contrasts are present at the boundaries of the illuminated circular window which results from the maximum angle of refraction at 48–9° with the normal (Snell's window). Contrasts are fixed by the compound eye and appropriate turning of the body axis orients the daphnid in an upwards or an obliquely downwards direction. A predisposition for a positively or negatively phototactic orientation seems to be the result of a disturbed balance of the two oscillators governing normal swimming. Some investigators have tried to study DVM at a laboratory scale during a 24 h cycle. To imitate nature, properties of a natural water column, such as a large temperature gradient, were compressed into a few cm. With appropriate light intensity changes, vertical distributions looking like DVM were obtained. The results can be explained by phototactic reactions and the artificial nature of the compressed environmental factors but do not compare with DVM in the field. A mechanistic model of DVM based on phototaxis is presented. Both, primary and secondary phototaxis is considered an extension of normal swimming. Using the light intensity changes of dawn and the differential enhancement of kairomones and food concentrations, amplitudes of DVM could be simulated comparable to those in a lake. The most important adaptive significance of DVM is avoidance of visual predators such as juvenile fish. However, in the absence of fish kairomones, small-scale DVMs are often present, which were probably evolved for UV-protection, and are realized by not enhanced phototaxis. In addition, the ‘decision-making mechanism’ was probably evolved as based on the enhanced phototactic reaction to accelerations in the rate of relative changes in light intensity and the presence of fish kairomones.     

Females compensate for moult‐associated male nest desertion in Hooded Warblers

Uniparental offspring desertion occurs in a wide variety of avian taxa and usually reflects sexual conflict over parental care. In many species, desertion yields immediate reproductive benefits for deserters if they can re‐mate and breed again during the same nesting season; in such cases desertion may be selectively advantageous even if it significantly reduces the fitness of the current brood. However, in many other species, parents desert late‐season offspring when opportunities to re‐nest are absent. In these cases, any reproductive benefits of desertion are delayed, and desertion is unlikely to be advantageous unless the deserted parent can compensate for the loss of its partner and minimize costs to the current brood. We tested this parental compensation hypothesis in Hooded Warblers Setophaga citrina, a species in which males regularly desert late‐season nestlings and fledglings during moult. Females from deserted nests effectively doubled their provisioning efforts, and nestlings from deserted nests received just as much food, gained mass at the same rate, and were no more likely to die from either complete nest predation or brood reduction as young from biparental nests. The female provisioning response, however, was significantly related to nestling age; females undercompensated for male desertion when the nestlings were young, but overcompensated as nestlings approached fledging age, probably because of time constraints that brooding imposed on females with young nestlings. Overall, our results indicate that female Hooded Warblers completely compensate for male moult‐associated nest desertion, and that deserting males pay no reproductive cost for desertion, at least up to the point of fledging. Along with other studies, our findings support the general conclusion that late‐season offspring desertion is likely to evolve only when parental compensation by the deserted partner can minimize costs to the current brood.     

Development of stress response in nestling pied flycatchers

Birds respond to unpredictable events by secreting corticosterone, which induces various responses to cope with stressful situations. However, the evidence is still elusive whether altricial nestlings perceive and respond to external stressors. We investigated the development of adrenocortical stress response to handling-related stressor in nestlings of a small passerine bird, the pied flycatcher (Ficedula hypoleuca). Nestlings were held in isolation from their parents during the experiment to ensure that they indeed respond to handling, not to parental alarm calls. We found that both 9- and 13-day-old nestlings were able to elicit hormonal stress response. Although baseline as well as stress-induced corticosterone levels rose slightly with age, the magnitude of difference between the control and stress-induced levels remained similar in both age groups. However, comparison with adults showed that the stress response of nestlings prior to fledging was still incomplete and significantly lower than in adults. Overall, our results indicate that altricial nestlings do respond to acute stressors, but on the contrary to previous predictions the development of corticosterone stress response during growth period is not gradual and varies remarkably between different passerine species.     

The effect of temperature on phototaxis and geotaxis by larvae of the crab Rhithropanopeus harrisi (Gould)

Investigations of the effect of sudden temperature change on the phototaxis of Stage I and IV zoeae upon stimulation from horizontal and vertical directions with 500-nm light indicate a temperature-induced geotactic response in larvae of the crab Rhithropanopeus harrisi (Gould). For the horizontal tests both zoea stages were reared at 20 °C. Stage I showed positive phototaxis at temperatures between 15 ° and 35 °C, while Stage IV responded over the range of 10–30 °C. For the vertical tests, larvae, reared at 25 °C, were stimulated with overhead lights. Stage I zoeae ascended at 15 °, 20 ° and 25 °C and descended at 5 °, 10 °, 30 ° and 35 °C. Stage IV zoeae ascended at 20 ° and 25 °C and descended at 5 °, 10 °, 15 °, 30 ° and 35 °C. Although the descent at high temperatures could result from a negative phototaxis, a reversal in phototactic sign at high temperatures was not found in the horizontal experiments and the same vertical movement pattern is observed in total darkness. Upon exposure to high temperatures near the water surface, larvae would descend by means of a positive geotaxis rather than a negative phototaxis. This response involves active swimming by Stage IV larvae and passive sinking by Stage I.     

Phototropin Influence on Eyespot Development and Regulation of Phototactic Behavior in Chlamydomonas reinhardtii

The eyespot of Chlamydomonas reinhardtii is a light-sensitive organelle important for phototactic orientation of the alga. Here, we found that eyespot size is strain specific and downregulated in light. In a strain in which the blue light photoreceptor phototropin was deleted by homologous recombination, the light regulation of the eyespot size was affected. We restored this dysfunction in different phototropin complementation experiments. Complementation with the phototropin kinase fragment reduced the eyespot size, independent of light. Interestingly, overexpression of the N-terminal light, oxygen or voltage sensing domains (LOV1+LOV2) alone also affected eyespot size and phototaxis, suggesting that aside from activation of the kinase domain, they fulfill an independent signaling function in the cell. Moreover, phototropin is involved in adjusting the level of channelrhodopsin-1, the dominant primary receptor for phototaxis within the eyespot. Both the level of channelrhodopsin-1 at the onset of illumination and its steady state level during the light period are downregulated by phototropin, whereas the level of channelrhodopsin-2 is not significantly altered. Furthermore, a light intensity–dependent formation of a C-terminal truncated phototropin form was observed. We propose that phototropin is a light regulator of phototaxis that desensitizes the eyespot when blue light intensities increase.     

Diel patterns of predation and fledging at nests of four species of grassland songbirds

Although it is common for nestlings to exhibit a strong bias for fledging in the morning, the mechanisms underlying this behavior are not well understood. Avoiding predation risk has been proposed as a likely mechanism by a number of researchers. We used video surveillance records from studies of grassland birds nesting in North Dakota, Minnesota, and Wisconsin to determine the diel pattern of nest predation and fledging patterns of four ground‐nesting obligate grassland passerines (Grasshopper Sparrow (Ammodramus savannarum), Savannah Sparrow (Passerculus sandwichensis), Bobolink (Dolichonyx oryzivorus), and Eastern Meadowlark (Sturnella magna)). We used the nest predation pattern as a surrogate for predation activity to test whether nestlings minimized predation risk by avoiding fledging when predation activity was high and preferentially fledging when predation risk was low. Predation activity was significantly lower starting 3 hr before sunrise and ending 3 hr after sunrise, followed by a transition to a period of significantly higher activity lasting for 4 hr, before declining to an average activity level for the rest of the diel period. There was little evidence that the four grassland bird species avoided fledging during the high‐risk period and Savannah Sparrow fledged at higher rates during that period. All four species had hours during the low‐risk period where they fledged at higher rates, but only Grasshopper Sparrow fledged preferentially during that period. Bobolink and Eastern Meadowlark had multiple hours with high fledging rates throughout the daytime period, resulting in no relationship between probability of fledging and predation risk. Given the species variability in fledging pattern seen in our study, it is unlikely that there is a universal response to any driver that affects time of fledging. Further study is needed to understand the complex interplay between species ecology and drivers such as physiology, energetics, and predation in affecting grassland bird fledging behavior.     

昆虫趋光性及其机理的研究进展

依据目前已有资料 ,从行为学、生理学及田间应用调查等方面概述了近几十年昆虫趋光性的国内外研究进展。其中 ,行为学与生理学研究较多 ,且两者结果基本一致 ,相互补充 ,为趋光性机制的假说提供了可靠的依据。关于昆虫趋光性机制的假说较多 ,其中报道较多的是光干扰假说、光定向行为假说和生物天线假说 3种 ,现在较为普遍接受的是前两者。光干扰假说是指刺眼作用干扰昆虫的正常活动导致趋光 ,而光定向行为假说则指昆虫趋光是由于光定向行为所致