An analysis of the spatial mechanisms responsible for electrosensitivity in rays-modelling results

Mathematical methods have helped to reveal the possible mechanisms of information conversion processes by the Lorenzinian ampullae during spatial detection of the dipole electric field. A stationary voltage distribution in the inhomogeneous media has been calculated by means of numerical methods. The skate body has been represented by a thin disk. If the dipole axis was positioned in the disk plane, potential distortions near the disk were negligibly small. In other cases the electrical field energy absorbed by ampullary groups has been dramatically reduced. It was supposed that body tissues served as the space filter of the electroreceptive system. Based on the result presented here, a new mechanism of the electrolocation was proposed. The ampullary groups were predicted to play the role of arrays which were able to perceive spatial characteristics of the external electric field. The theoretical analysis of ampullary groups circular sensitivity diagrams supported the prediction.     

Electrostatics and electrodynamics of bacteriorhodopsin.

The stationary electric dichroism of bacteriorhodopsin is in qualitative, but not quantitative, agreement with the orientation function for disks having a permanent dipole directed perpendicular to the plane and an induced dipole in the plane. Fits of the orientation function to data measured at low field strengths demonstrate: an increase of the permanent dipole moment mu with the square of the disk radius r2, whereas the polarizability alpha increases with r4; the ionic strength dependence is small for mu and clearly stronger for alpha; the permanent dipole moment is 4x10(6) D at r = 0.5 micron. According to the risetime constants, the induced dipole does not saturate and increases to 4x10(8) D at 40 kV/cm and r = 0.5 micron. The data indicate that the permanent dipole is not of some interfacial character but is due to a real assymetry of the charge distribution. The experimental dipole moment per protein monomer is approximately 55 D, whereas calculations based on the structure of Grigorieff et al. (Grigorieff, N., T.A. Ceska, K.H. Downing, J.M. Baldwin, and R. Henderson. 1996. Electron-crystallographic refinement of the structure of bacteriorhodopsin. J. Mol. Biol. 259:393-421) provide a dipole moment of approximately 570 D. The difference is probably due to a nonsymmetric distribution of charged lipid residues. It is concluded that experimental dipole moments reflect the mu-potential at the plane of shear for rotational diffusion, in analogy to the sigma-potential used for translational diffusion. It is suggested that the permanent dipole of bacteriorhodopsin supports proton transport by attraction of protons inside and repulsion of protons outside of the cell. Dichroism rise curves at field strengths between E = 150 and 800 V/cm reveal an exponential component with time constants tau 3r in the range between 1 and 40 ms, which is not found in Brownian dynamics simulations on a disk structure using hydrodynamic and electric parameters characteristic of bacteriorhodopsin disks. The experimental data suggest that this process reflects a cooperative change of the bacteriorhodopsin structure, which is induced already at a remarkably low field strength of approximately 150 V/cm.     

Theoretical analysis of pre-receptor image conditioning in weakly electric fish

Electroreceptive fish detect nearby objects by processing the information contained in the pattern of electric currents through the skin. The distribution of local transepidermal voltage or current density on the sensory surface of the fish's skin is the electric image of the surrounding environment. This article reports a model study of the quantitative effect of the conductance of the internal tissues and the skin on electric image generation in Gnathonemus petersii (Günther 1862). Using realistic modelling, we calculated the electric image of a metal object on a simulated fish having different combinations of internal tissues and skin conductances. An object perturbs an electric field as if it were a distribution of electric sources. The equivalent distribution of electric sources is referred to as an object's imprimence. The high conductivity of the fish body lowers the load resistance of a given object's imprimence, increasing the electric image. It also funnels the current generated by the electric organ in such a way that the field and the imprimence of objects in the vicinity of the rostral electric fovea are enhanced. Regarding skin conductance, our results show that the actual value is in the optimal range for transcutaneous voltage modulation by nearby objects. This result suggests that "voltage" is the answer to the long-standing question as to whether current or voltage is the effective stimulus for electroreceptors. Our analysis shows that the fish body should be conceived as an object that interacts with nearby objects, conditioning the electric image. The concept of imprimence can be extended to other sensory systems, facilitating the identification of features common to different perceptual systems.     

Mechanism of perception of changes in the magnetic field by ampullae of lorenzini of elasmobranchs

The mechanisms of reception of changes in the magnetic field by electroreceptor formations were investigated in experiments on Black Sea rays in which spike activity was recorded from single nerve fibers connected with the ampullae of Lorenzini. The responses of the ampullae of Lorenzini to magnetic stimulation were shown to be due to induced electric currents creating potential gradients in the body tissues of the fish and the sea water. On the basis of differences in responses of different ampullae to magnetic stimulation, it is possible to distinguish between magnetic stimuli and other stimuli acting on the electroreceptor system. Potentiation of the receptor response to magnetic stimulation was found as the fish came closer to the "shore." The mechanisms and biological importance of reception of changes in the magnetic field by the ampullae of Lorenzini are discussed.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 10, No. 1, 75–83, January–February, 1978.     

Functional characteristics of electroreceptors of the Turkestan catfish

Acute experiments to record spike activity from single fibers of the lateral line nerve of the Turkestan catfish revealed electroreceptor formations which, in their functional characteristics, were similar to the ampullated electroreceptors of other freshwater fish (the so-called small pit organs). The threshold intensity of the uniform electric field was 1 µV/cm. A voltage drop on the skin of the fish was shown to be an effective stimulus for the electroreceptors. A spike response to a change in the magnetic field was found for the first time in electroreceptors of freshwater fish. The threshold level of magnetic induction for a velocity of rotation of the permanent magnet of 1 m/sec was 2.9 · 10^–4 T. Temperature and mechanical sensitivity of the electroreceptors was determined. The biological significance of electroreceptors of the Turkestan catfish is discussed.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. M. I. Kalinin Andizhan Medical Institute, Ministry of Health of the Uzbek SSR. S. M. Kirov Murmansk Marine Biological Institute, Kola Branch, Academy of Sciences of the USSR, Dal'nie Zelentsy. Translated from Neirofiziologiya, Vol. 12, No. 5, pp. 508–516, September–October, 1980.     

Automatic Realistic Real Time Stimulation/Recording in Weakly Electric Fish: Long Time Behavior Characterization in Freely Swimming Fish and Stimuli Discrimination

Weakly electric fish are unique model systems in neuroethology, that allow experimentalists to non-invasively, access, central nervous system generated spatio-temporal electric patterns of pulses with roles in at least 2 complex and incompletely understood abilities: electrocommunication and electrolocation. Pulse-type electric fish alter their inter pulse intervals (IPIs) according to different behavioral contexts as aggression, hiding and mating. Nevertheless, only a few behavioral studies comparing the influence of different stimuli IPIs in the fish electric response have been conducted. We developed an apparatus that allows real time automatic realistic stimulation and simultaneous recording of electric pulses in freely moving Gymnotus carapo for several days. We detected and recorded pulse timestamps independently of the fish’s position for days. A stimulus fish was mimicked by a dipole electrode that reproduced the voltage time series of real conspecific according to previously recorded timestamp sequences. We characterized fish behavior and the eletrocommunication in 2 conditions: stimulated by IPIs pre-recorded from other fish and random IPI ones. All stimuli pulses had the exact Gymontus carapo waveform. All fish presented a surprisingly long transient exploratory behavior (more than 8 h) when exposed to a new environment in the absence of electrical stimuli. Further, we also show that fish are able to discriminate between real and random stimuli distributions by changing several characteristics of their IPI distribution.     

Growth and fecundity of Daphnia after diapause and their impact on the development of a population

Laboratory and field investigations revealed that the life history traits of exephippial and parthenogenetic generations of Daphnia differ substantially. Daphniids hatching from resting eggs grow faster and their definitive body sizes are bigger than of hatchlings from subitaneous eggs. Size at maturity for exephippial animals is significantly larger. In spite of this, they mature a few days earlier than parthenogenetic females. In this study, the difference was 3–4 days for the laboratory experiments and 1–3 days for the field. Fecundity of the exephippial generation is markedly higher. Here, the clutch size for this generation was up to 3.5–4.0 times as large as for the parthenogenetic generation. Moreover, obtained results suggest that the relationship between clutch size and body length for both generations differ significantly.Estimates of the intrinsic rate of increase for field Daphnia populations demonstrated that life history traits of exephippial animals lead to a two or threefold higher rate of increase in the conditions of invertebrate predation pressure. Under moderate fish pressure, obtained r values for the daphniids hatching from resting eggs were larger than those from subitaneous. High growth rate of exephippial females is disadvantageous only under the conditions of severe pressure by fish. Obtained results suggest that hatchlings from diapausing eggs an acceleration of population increase by several times during the beginning of the development of a population with periodical re-establishment from resting eggs.     

Effect of body orientation in the gravitational field on directional sensitivity of the cercal system of neurons in crickets

Changes in the spatial orientation of three-dimensional directional sensitivity diagrams of neurons of the terminal abdominal ganglion of the cricket during body tilting were studied. Spike responses were recorded from neurons of the ganglion to acoustic stimuli in different directions, with the cricket's body tilted at different angles to the horizontal plane. During tilting of the cricket's body the orientation of the directional sensitivity diagrams was found to change parallel with the orientation of the body. Neurons of the abdominal ganglion are excited by cercal sensillae, among which there are receptors which respond to changes in the position of the cricket's body in the gravitational field (gravity receptors). The results suggest that cercal gravity receptors have no specific influence on the directional sensitivity of neurons of the first central division of the cercal system.Institute for Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 12, No. 6, pp. 604–611, November–December, 1980.     

Molecular organization of bacteriochlorophyll in chlorosomes of the green photosynthetic bacteriumChloroflexus aurantiacus: Studies of fluorescence depolarization accompanied by energy transfer processes

Examination was made of changes in fluorescence polarization plane by energy transfer in the chlorosomes of the green photosynthetic bacterium,Chloroflexus aurantiacus. Fluorescence anisotropy in the picosecond (ps) time region was analyzed using chlorosomes suspended in solution as well as those oriented in a polyacrylamide gel. When the main component of BChlc was preferentially excited, the decay of fluorescence anisotropy was found to depend on wavelength. In the chlorosome suspension, the anisotropy ratio of BChlc changed from 0.31 to 0.24 within 100 ps following excitation. In the baseplate BChla region, this ratio decreased to a negative value (–0.09) from the initial 0.14. In oriented samples, the degree of polarization remained at 0.68 for BChlc, and changed from 0.25 to –0.40 for the baseplate BChla by excitation light whose electric vector was parallel to the longest axis of chlorosomes. In the latter case, there was a shift from 0.30 to –0.55 by excitation perpendicular to the longest axis. Time-resolved fluorescence polarization spectra clearly indicated extensive changes in polarization plane accompanied by energy transfer. The directions of polarization plane of emission from oriented samples were mostly dependent on chlorosome orientation in the gel but not on that of the polarization plane of excitation light. Orientations of the dipole moment of fluorescence components was consistent with that of absorption components as determined by the linear dichroism (Matsuura et al. (1993) Photochem. Photobiol. 57: 92–97). A model for molecular organization of BChlc anda in chlorosomes is proposed based on anisotropic optical properties.     

Intertidal territoriality and time-budget of the surgeonfish, Acanthurus lineatus, in American Samoa

Synopsis The herbivorous surgeonfish Acanthurus lineatus aggressively maintained feeding territories in the surf zone of the outer reef flat in American Samoa. Intertidal territories were re-established each morning, as well as after displacement by low tides or rough surf. Day-to-day site fidelity of recognizable individuals was high: 99.9% return rate per day for adults (15–20 cm), 99.6% for juveniles (8–13 cm), and 97.2% for recruits (2.5–5 cm). Fish fed on turf algae primarily in the afternoon (80% of available time), and spent 10% of their time on active territorial defense and 2–13% of their time on forays from their territory. On average, a fish defended its territory 1900 times daily and took 17 000 bites (= 7400 bites m^–2 d^–1), but rough surf reduced feeding by 60% and defense by 75%. High territorial defense requirements significantly reduced feeding rates. Although the distribution and behavior of this species in Samoa was in large part similar to that reported for it elsewhere (Australia, Indian Ocean), there were notable differences: in Samoa A. lineatus densities within colonies were greater (0.4 fish m^–2), territory size was smaller (2.3 m²), and defense rate against intruders was greater (2.5 attacks min^–1). These differences in Samoa may be related to their smaller body size, greater abundance or increased food supply caused by hurricane damage to reefs which has enhanced the algal turfs that A. lineatus feeds upon.     

Scanning proton microprobe analysis of strontium in an arctic charr,Salvelinus alpinus,otolith: implications for the interpretation of anadromy

Synopsis Scanning proton microprobe imaging has been used to map the distribution of strontium (Sr) in an arctic charr, Salvelinus alpinus, otolith. X-ray maps show that Sr variation follows an oscillatory zoned pattern which correlates with optically resolved zones (annuli). The internal region of the otolith shows optical zonation but no significant Sr content. Comparison of the optical image with the Sr X-ray map provides evidence of when the fish entered a high Sr environment. This may be interpreted as when the fish migrated from fresh water to a marine environment; this particular fish first migrated to sea in its eighth year. Micro-PIXE point analysis was used to determine the abundance of Sr in annuli along a radius from the nucleus to an outer edge; Sr levels in the internal region were about 10 ppm and variation in the outer zoned region was between 62 and 175 ppm. Detection limits for Sr are in the 1–2 ppm range. Other trace elements, such as transition elements, were found to be present in some growth regions in the 1–20 ppm range. Proton beam analysis is a non-destructive analytical technique capable of preserving the spatial integrity of trace-element data in otoliths such that element distribution may be linked to the growth structure of the otolith.     

Characteristics of reaction field and the reactive distance of a planktivore, Pseudorasbora parva (Cyprinidae), in various environmental conditions

The characteristics of the reaction field and the reactive distance of the Stone moroko (Pseudorasbora parva) were studied under three environmental conditions (structural complexity, light intensity and turbidity) and three prey sizes. In optimal experimental conditions, under no structural complexity, light intensity of 200 lux and turbidity less than 1 NTU (Nephelometric Turbidity Units), the cross-section of the reaction field was found to be elliptic with a bearing angle larger than the elevation angle, but both angles changed slightly depending on environmental conditions. The reactive distance was large, and the fish frequently attacked prey that was located within 15–60 degrees to each side from the frontal direction of a fish (i.e., ± 15 degrees from the axis of the fish body) horizontally. In the light intensity below 50 lux or turbidity above 10 NTU, however, the attack frequency and the reactive distance in the frontal direction of a fish did not differ from other horizontal directions in the reaction field. The average reactive distance increased proportionally with increasing strand distance, but it gradually reached a constant value for strand distances greater than about 3.6 times the fish body length. The average reactive distance increased in the light intensity range of 10–200 lux and decreased negatively with turbidity increasing. The average reactive distance increased with larger prey size, but the rate of increase of the reactive distance gradually decreased.     

Voltage-dependent channel formation by rods of helical polypeptides

Summary The voltage-dependence of channel formation by alamethicin and its natural analogues can be described by a dipole flip-flop gating model, based on electric field-induced transbilayer orientational movements of single molecules. These field-induced changes in orientation result from the large permanent dipole moment of alamethicin, which adopts -helical conformation in hydrophobic medium. It was, therefore, supposed that the only structural requirement for voltage-dependent formation of alamethicin-type channels might be a rigid lipophilic helical segment of minimum length.In order to test this hypothesis we synthesized a family of lipophilic polypeptides—Boc-(Ala-Aib-Ala-Aib-Ala) _n -OMe,n=1–4—which adopt -helical conformation forn=2–4 and studied their interaction with planar lipid bilayers. Surprisingly, despite their large difference in chain length, all four polypeptides showed qualitatively similar behavior. At low field strength of the membrane electric field these polypeptides induce a significant, almost voltage-independent increase of the bilayer conductivity. At high field strength, however, a strongly voltage-dependent conductance increase occurs similar to that observed with alamethicin. It results from the opening of a multitude of ion translocating channels within the membrane phase.The steady-state voltage-dependent conductance depends on the 8^th–9^th power of polypeptide concentration and involves the transfer of 4–5 formal elementary charges. From the power dependences on polypeptide concentration and applied voltage of the time constants in voltage-jump current-relaxation experiments, it is concluded that channels could be formed from preexisting dodecamer aggregates by the simultaneous reorientation of six formal elementary charges. Channels exhibit large conductance values of several nS, which become larger towards shorter polypeptide chain length. A mean channel diameter of 19 Å is estimated corresponding roughly to the lumen diameter of a barrel comprised of 10 -helical staves. Similar to experiments with the N-terminal Boc-derivative of alamethicin we did not observe the burst sequence of nonintegral conductance steps typical of natural (N-terminal Ac-Aib)-alamethicin. Saturation in current/voltage curves as well as current inactivation in voltage-jump current-relaxation experiments are found. This may be understood by assuming that channels are generated as dodecamers but, while reaching the steady state, reduce their size to that of an octamer or nonamer. We conclude that the overall behavior of these synthetic polypeptides is very similar to that of alamethicin. They exhibit the same concentration and voltage-dependences but lack the stabilizing principle of resolved channel states characteristic of alamethicin.     

Studies on the age and growth of the bata,Labeo bata (Ham.) (Cyprinidae,Teleostei) from the river Kali,India

Age and growth of Labeo bata (Ham.) was studied by the analysis of annuli found on the scale and by length-frequency distribution. The fish attained lengths of 131, 194, 236, 277, 314, 341 and 364 mm at the end of the 1st, 2nd, 3rd, 4th, 5th, 6th and 7th years of life respectively. The increase in length of scale bears a constant relationship with the increase in length of fish, and regression analysis yielded a straight line between scale and body length. Calculated values could be expressed as: Y = –2.534 + 0.064 XThe growth rate of the fish was found high during the 1st and 2nd years and decreased gradually afterwards till the 7th year. Both sexes showed more a or less similar growth rate and attained a similar longevity. Growth trend of the fish confirmed the von Bertalanffy growth equation: Lt = 450(l–e^{–0.2165(t+0.5963))} The seasonal growth curve was chiefly influenced by feeding intensity in fishes of 1st year class while in adults it was affected by feeding intensity as well as by maturation of the gonads.     

The Development of Vision in the Zebrafish (Danio rerio)

We studied the development and maturation of the visual system by determining when zebrafish begin to see and to move their eyes. This information was correlated with the time courses of the development of the retina, the retinofugal projection, the retinal image, and the extraocular muscles, to obtain an integrated picture of early visual development. Two visual behaviors were monitored over 48–96 hr postfertilization (hpf). The startle response (body twitch) was evoked by an abrupt decrease in light intensity. The optokinetic response (tracking eye movements) was evoked by rotation of a striped drum. Visually evoked startle developed over 68–79 hpf, more than 20 hr after the onset of a touch-evoked startle. It was not seen in eyeless fish, excluding a role for nonretinal light senses. Tracking eye movements developed over 73–80 hpf. They were always in the direction of drum rotation, even when the fish had been light deprived from blastula stage, ruling out a “trial and error” period of learning to track the drum. The image formed by the ocular lens was examined in intact fish made transparent by suppressing the formation of melanin. The eye was initially far sighted and gradually improved, so that by 72 hpf the image plane coincided with the photoreceptor layer. The extraocular muscles assumed their adult configuration between 66 and 72 hpf. Thus, the retinal image and functional extraocular muscles appeared nearly simultaneously with the onset of tracking eye movements and probably represent the last events in the construction of this behavior.     

Ontogeny of vision in the Antarctic fish Pagothenia borchgrevinki (Nototheniidae)

Summary Pagothenia borchgrevinki ranging in size from 63 to 245 mm were captured from beneath sea ice in McMurdo Sound by fishing and diver collection. Changes in ocular morphology with increasing body size were measured, and assessed in relation to ingested prey. Relative eye size was highest among smaller fish (<100mm total length), and declined with increasing fish size. This was accompanied by a decrease in cone density in the retina from a maximum of 14,200 mm^–2 in the smallest fish examined (63 mm), to 1000 mm^–2 in a 220 mm long fish. Theoretical acuity was lowest among fish at either end of the size range examined (minimum separable angle 40–50) but approximately constant over the remainder of the size range (25–40). Rod density also decreased with increasing body size but rod numbers per unit visual arc were relatively constant, except in the smallest fish, where angular rod density was low. The same prey taxa occurred in fish of all sizes; however, prey items smaller than about 1.5 mm were not taken by fish of any body size.     

Functional properties of lamprey electroreceptors

Spike discharges in nerve fibers, evoked by stimulation of electroreceptors by an electric field directed along the body axis (square pulses of current and a sinusoidal current) were recorded in the lampreyLamperta fluviatilis (L.). Excitation of electroreceptors was shown to arise through the action of the cathode. Minimal values of electric field at which appreciable changes took place in spike activity were 10–60 µV/cm for different nerve fibers. The optimal frequency range of sinusoidal electrical stimulation was 0.05–0.5 Hz. It is suggested that electroreceptors of the Agnatha (caudata) and of gnathostomatous cartilaginous fish share a common origin.Murmansk Marine Biological Institute, Kola Branch, Academy of Sciences of the USSR, Dal'nie Zelentsy, Murmansk Region. Translated from Neirofizioloigya, Vol. 16, No. 1, pp. 105–110, January–February, 1984.     

云南绿汁江鱼类的形态特征及其适应意义

绿汁江的鱼类大致可分为侧扁型,圆筒型和平扁型三种类型。在各种类型中,鱼体上又有不同的形态结构产生。以形态－功能－适应三位一体分析法为指导原则,探讨和阐述了鱼类的形态结构与栖息环境的相互关系。     

Aspects of the fecundity of the feral catfish,Clarias macrocephalus (Gunther), population obtained from the rice fields used for rice-fish farming,in Malaysia

The catfish, Clarias macrocephalus, harvested from the irrigated rice fields of Malaysia, exhibited spawning activities that are influenced by water level management. The multiple spawning potential of the fish is negated by declining water level and the fish spawned only once per growing season. Although oocyte development started in conjunction with water level increase, subsequent oocyte development did not take place due to low water level. Absolute and relative fecundity also decreased with declining water level, however, mean relative fecundity is relatively constant (17.6–53.7 oocytes g^–1 body weight) and not related to body size. Absolute fecundity ranged between 785 to 14066 oocytes female^–1, whereas relative fecundity is between 7.1 to 83.5 oocytes g^–1 body weight, respectively.     

Assessment of Grey Heron predation on fish communities: the case of the largest European colony

The fish predation rate by Grey Heron Ardea cinerea was studied during two breeding seasons (1987–88) in the largest European colony at the Lake of Grand-Lieu (Loire-Atlantique, France). The herons' diet was compared to the available fish population of its main feeding area, the marsh of Bourgneuf (16000 ha) which is composed of former salt pans and meadows drained by a dense network of shallow ditches. This study is the first attempt to assess the predation exerted by an ardeid colony on a fish community over such an extensive natural environment. It also provides the first data about the abundance and the structure of fish communities in shallow coastal dyked marshes. For this purpose, two different sampling methods were used according to the water's salinity. In fresh waters, electrofishing was used as the removal method, and density estimates were calculated with Carle & Strub estimator (1978). Fish were caught in randomly selected stations (sections of ditches enclosed by two 5 mm mesh nets). In brackish waters, pools and ditches were drained. The distribution of the herons at the feeding areas was determined by direct observations, by counting flights from the colony, and by radio-tracking. The diet was investigated by observing adult herons on the foraging areas, and by analyzing the prey regurgitations of the young at the nests. The global food consumption was assessed from Marion (1988), according to the birds' activity determined during 5 years of radio-tracking. Altogether, at least 39 species of fish were available in the herons' feeding area (during the reference period, 87–88) and the mean fish biomass was 270 kg per ha of open water, or 30 kg per ha of marsh (open water = 11.2% of the marsh area). The fish community was dominated by eel Anguilla anguilla (145 kg ha^–1, 50,8% of the total biomass), and catfish Ictalurus melas (40 kg ha^–1, 14%). Except for small and inaccessible species (living in the deepest parts of the marsh), heron diet was very similar to fish species composition of the community occurring in the marsh. The catfish was the species captured most frequently by the heron (45% of the mass), the eel was second with 28% of the mass. The catfish was probably over represented in the diet considering that they are caught in catfish-dumps created by professional fishermen at Grand-Lieu lake, in order to reduce the density of this undesirable species. Inversely, small species such as Gasterosteus aculeatus were not found in the diet whereas they are very numerous in the marsh. On average herons of Grand-Lieu colony catch 1.92 kg of fish per ha of marsh (6% of the fish standing crops in the marsh) during the breeding season, the main predation period.