Vertical distribution and diel migration of rotifers in a Parana River floodplain lake

The vertical distribution of zooplankton rotifers in the open waters of Laguna El Tigre was investigated. Rotifers showed a relatively uniform distribution throughout the water column. This pattern of distribution was maintained during the year and did not show variations in relation to hydrologic phases of inundation and isolation of the lake. Diel vertical migration of rotifers from the limnetic and the littoral area was investigated too. In littoral area rotifers exhibited a reverse migration, whereas in the limnetic the movements were less conspicuous. Horizontal migration was observed too, and there were interactions between horizontal and vertical distribution. Predation and competition offer a possible explanation.     

Seasonal variation in the diel vertical distribution of the migratory alga Cryptomonas marssonii (Cryptophyceae) in a small,highly humic lake

The diel vertical distribution patterns of a migratory alga Cryptomonas marssonii in a small, steeply stratified humic lake were investigated during a summer season (10 diurnal experiments between May and September) using a close-interval Blakar-type sampler. The results indicate that the cells were phototactic; they were typically concentrated at the surface or subsurface during daylight, while in darkness the highest densities were recorded in deeper water, usually near the upper limit of anoxia. During a dense blue-green bloom in August the cells of C. marssonii were also concentrated by day into the same water layer, where oxygen was depleted. However, the cells seemed to avoid totally anoxic water. Because the vertical distribution pattern of C. marssonii had special diurnal and seasonal characteristics, care is needed when designing a sampling programme for a phytoplankton population dominated by this species.     

Seasonal variations in the diel vertical distribution of phytoplankton and zooplankton in a shallow pond

The seasonal succession of phytoplankton diversity, and the variations in the diel vertical distribution of phyto‐ and zooplankton were investigated in a small shallow pond (1.7 m water depth) in 2003. It was inferred that the water tended to stratify weakly in the daytime from February to June. In February and April, the green alga Golenkinia radiata Chodat dominated the phytoplankton assemblage. The cell density of G. radiata greatly decreased in April, when rotifers increased near the bottom. The vertical mixing was attenuated in June, large populations of the euglenoids (Lepocinclis salina Fritsch, Phacus acuminatus Stokes, Trachelomonas hispida (Perty) Stein et Deflandre) developed, and the cyanobacterium Aphanizomenon flos‐aquae var. klebahnii Elenk. appeared at low density. Euglenoids and A. flos‐aquae were mostly distributed in the bottom layer. In late September, when the water was mixed throughout the day, euglenoids and A. flos‐aquae were distributed evenly throughout the water column. The zooplankton (cyclopoid copepods and rotifers) densities in September were the lowest throughout the year. The vertical mixing increased in November, and the phytoplankton community was composed of A. flos‐aquae, P. acuminatus, T. hispida and the green alga Ankistrodesmus falcatus (Corda) Ralfs. In November, at the final stage of water bloom of A. flos‐aquae, its population density decreased with depth. The two euglenoids exhibited similar cell distributions at 0.8 m and 1.6 m during 1–3 November. A. falcatus was distributed evenly throughout the water column; however, when the vertical mixing lessened, the cells at the surface started to sink. Copepod nauplii and rotifers appeared at high densities in November. Seasonal variation in the phytoplankton community structure in the pond seemed to be related to the vertical mixing of the water. In addition, zooplankton, especially rotifers, might play an important role in initiating a spring clear‐water phase and in the bloom collapse of A. flos‐aquae.     

浮游动物昼夜垂直迁移机理的主要假说及其研究进展

对有关浮游动物昼夜垂直迁移(DVM)机理的实验、假说以及理论模型方面的研究进.展进行了综述。昼夜垂直迁移通常指常规迁移(傍晚上升,拂晓下降),其行为过程不仅影响浮游动物的垂直分布,而且也间接地影响其水平分布及生活史特征,对浮游动物在一定水域的种群维持和补充具有重要的意义。与垂直迁移机理有关的假说主要有光驱动假说、逃避捕食者假说、能量和资源利用假说等,其中逃避捕食者假说已得到大量的实验证实。其理论模型研究一般在两个时间尺度上进行,一个是短期的行为机制研究,另一个从长期的生活史策略的角度考虑。随着实验技术的进步和理论及建模工具的发展,这两个时间尺度的研究正逐渐统一到一个框架下进行。     

Vertical distribution of phytoplankton in Lake Xolotlán (Managua)

A study of the vertical distribution of phytoplankton was carried out at the end of the rainy season (October–November), when wind activity is lower and the probability for phytoplankton stratification is enhanced. Samples were taken every other day at 0, 0.5, 1.0 m and, thereafter, for every meter up to the bottom. The number of individuals and their biomass were determined for each sampling depth and expressed as wet weight and chlorophyll-a, respectively. The results indicate that there were no significant differences (ANOVA one way, P<0.05) with regard to the vertical distribution of phytoplankton biomass over the 15 m of the water column. This result reflects the polymictic condition of Lake Xolotlán (Managua).     

The influence of the diel climatic cycle on the depth-time distribution of phytoplankton and photosynthesis in a shallow equatorial lake (Lake Baringo,Kenya)

Lake Baringo is a shallow equatorial lake. This paper reports a diel study of the depth-time distribution of phytoplankton and photosynthesis at one location in Lake Baringo on 10 March 1989. The water column shows a pattern of diurnal stratification probably accentuated by the high turbidity of the water and therefore rapid attenuation of solar energy. This stratified pattern breaks down at night due to atmospheric cooling and the regular onset of winds in the early evening. The phytoplankton is dominated byMicrocystis aeruginosa with some associated epiphytes. It concentrates in the narrow euphotic zone during the diurnal period of stratification due to buoyancy of theMicrocystis; evening breakdown of the thermocline results in the phytoplankton being mixed throughout the water column. A series of measurements of photosynthesis throughout the diurnal period gives an areal rate of 3.8 g O₂ m⁻² d⁻¹. The relationship between this value and the level of fish exploitation in Lake Baringo is discussed. The diel cycle in Lake Baringo is interpreted as dominating over any seasonal limnological cycle in the lake.     

Diurnal changes in the vertical distribution of cladocerans in a biomanipulated lake

The diel vertical migration of cladoceran zooplankton in LakeVesijärvi was studied after a distinct improvement of the waterquality following mass removal of fish. Four out of seven speciesshowed diurnal changes in vertical distribution. The two speciesdominating at 0–10 m, Bosmina crassicornis and Daphniacucullata, migrated reverse, while hypolimnetic Bosminalongispina and Ceriodaphnia quadrangula ascended at night.The migration did not cross the thermocline, suggesting thatzooplankton had a minor role in restricting the availability ofphosphorus for phytoplankton. The reverse migration in theepilimnion was likely connected to the feeding behaviour of thedominating planktivores. During night, smelt (Osmeruseperlanus (L.)) migrated from the hypolimnion to the surface tofeed on daphnids, while roach (Rutilus rutilus (L.)) thatinhabited the 5–15 m depth, was day-active and fed mainly on Bosmina. In the hypolimnion, the cladocerans used the low oxygenconcentration as a refuge and at night migrated to layers moreprofitable for feeding. This suggests that the predation by fishwas the ultimate factor of the vertical migration, but otherenvironmental factors determined its magnitude.     

Quantification of diel vertical migration by micronektonic taxa in the northeast Atlantic

Analyses of day/night changes in the bathymetric distribution of micronektonic biomass at 16 stations in the northeastern Atlantic, sampled between 1978 and 1994, provided quantitative estimates of the organic carbon fluxes associated with diel vertical migration of individual taxa of micronekton. Gelatinous taxa contributed 50–80% of the integrated standing crop by volume but, apart from tunicates, contributed relatively little to the active migratory fluxes when expressed in terms of carbon. Total micronektonic migratory fluxes into the upper 200 m ranged from 12.5 to 58 mgC per m². At 15 stations, fish and pteropods provided 50–80% of the fluxes into the upper 100, 200 and 400 m. At one station, tunicates (pyrosomes) contributed substantially. Wherever tunicates or the medusa Pelagia were swarming, migrations by other taxa appear to be suppressed. The mean proportions of the stock (in terms of biomass) of each of the dominant migratory taxa entering and leaving the upper 100 m were 23% for tunicates, 18% for fish, 22% for pteropods, 8% for decapod crustaceans and 23% for euphausiids. The maximum proportions for these five taxa were 90%, 60%, 75%, 25% and 75%, respectively. Similar estimates of the mean fluxes into and out of the upper 400 m were generally higher: 19% for tunicates, 39% for fish, 28% for pteropods, 49% for decapods and 55% for euphausiids; the respective maxima were 99%, 74%, 99%, 72% and 91%. It is estimated for fish that if these migrations occur throughout the year, they will result in an active carbon export (both POC and DOC) from the wind-mixed layer and immediate sub-thermocline depths of about 500% of the mean annual standing stock. If this estimate can be extended to other taxa, then the material fluxes resulting from these active migrations will be quantitatively similar to those resulting from the deposition of phytodetritus at temperate latitudes.     

Acoustic observation of diel vertical migration and shoaling behaviour in Atlantic redfishes

Acoustic methods were used to study Atlantic redfishes Sebastes spp. vertical migration and shoaling behaviour in Newfoundland waters. Redfishes exhibited consistent patterns of vertical migration in winter, spring and summer, but pelagic shoals were not observed in winter. Pelagic daytime aggregations were generally in close proximity to dense patches of redfishes along the sea floor. Pelagic shoals exhibited high degrees of variability in size, shape and density. Attempts to explain variations in shoal density and area with features of shoal position and structure were unsuccessful. Nearest neighbour distance between fish in shoals had a lower limit near one body length. During the night, fishes were dispersed in the water column and distributions were more homogenous. Diel vertical migration appeared to be a foraging strategy, in which redfishes followed the migration of their euphausiid prey.     

Distribution and diel migration of macroinvertebrates within dense submerged vegetation

1. We studied vertical and horizontal distribution of macroinvertebrates within a dense stand of Chara spp. in Lake Krankesjön, southern Sweden. Invertebrates were sampled at three depths within the vegetation and at three distances from the vegetation edge during day and night in July and August. Corresponding samples of oxygen content of the water were taken.
2. The densities (number of invertebrates per unit plant biomass) of most invertebrate taxa were generally lower in the upper layers of the vegetation than in the layers close to the sediment. The densities of several taxa ( Asellus aquaticus , Cloëon sp. and Polycentropodidae), as well as total density of invertebrates, were higher at the edge than in the innermost parts of the macrophyte stand, whereas snail densities generally were highest at the innermost sites.
3. Densities of A. aquaticus , Chironomidae larvae and Helobdella stagnalis generally increased at night. These taxa appear to undertake a diel vertical migration within the vegetation, towards, or even down to the sediment in daytime and up into the vegetation, in some cases to the vegetation surface, at night. Factors underlying the diel vertical migration are discussed, as are their ecological consequences.     

On the combined analysis of proximate and ultimate aspects in diel vertical migration (DVM) research

Although evolutionary ecologists agree that proximate and ultimate aspects are two sides of one coin, they are seldom interested in studies on physiological and behavioural mechanisms at the base of ecological phenomena. Nevertheless, these mechanisms are objects of selection and evolved to realise adaptive significances. This paper is a plea to bring both fields closer together, and, by means of an example of Diel Vertical Migration of Daphnia, some proximate and ultimate aspects are discussed. It is argued that light changes, not fish kairomone, is the primary cause for an individual to swim downwards at dawn and upwards at dusk. However, what is called a causal factor might differ when ecosystems or individuals are studied. In addition, causality in ecology is not simple, and has the character of a `set of necessary conditions'. To illustrate the importance of proximate analyses in DVM, two basic response mechanisms are discussed: Photobehaviour system 1 and 2. The physiological character of these systems leads to a fixed type of migration or to a phenotypically induced DVM, respectively. The adaptive significance of the first might be a reduction of the hazardous effects of UV radiation and of the second a lowering of mortality due to visually hunting predators.     

Vertical, lateral and longitudinal movement of zooplankton in a large river

1. The spatial distribution and movement patterns of zooplankton in large rivers are little known compared with those in lake environments. We conducted a series of studies in the Ohio River (U.S.A.) during the low flow period to assess diel vertical (DVM), longitudinal and lateral movement of crustacean zooplankton. 2. The dominant large zooplankter, the copepod Eurytemora affinis, showed a consistent vertical migration pattern of daytime ascent and night‐time descent during all sampling periods – the reverse of the most common migratory pattern of zooplankton in lakes. The cladoceran Bosmina migrated in a similar way in two of the three sampling periods. Surveys taken longitudinally in the river showed similar trends for both taxa. 3. During the lateral surveys, E. affinis was significantly more abundant in the shallow littoral zone during the night than in the daytime. The combination of vertical and lateral movement patterns along with the diel distribution of zooplanktivorous fish suggest that these movements are a predator‐avoidance mechanism. 4. Sampling programmes in large rivers should consider that larger zooplankton such as E. affinis may not be randomly distributed in the river channel and behaviours such as diel vertical migration may be just as evident in river habitats as in lakes.     

Thermal stability and phytoplankton distribution

Thermal stability is the potential of water columns to mix, and has long been known to fundamentally influence the vertical and temporal distribution of phytoplankton. Essentially this is because it indirectly controls the amount of light available to phytoplankton.Under stable conditions of strong temperature gradients algal species (or assemblages of associated species) distribute vertically because they have sufficient time to exploit the attenuated light field at their preferred depths. This encourages a species diversity which, in the Southern Hemisphere, is especially exemplified by the extremely stable conditions under the permanent ice of Antarctic lakes.In other lakes stability commonly encourages growth of blue-green algae by permitting their positive buoyancy to place them in optimal light conditions, and by inhibiting the resuspension of competing non-buoyant species. Analogous patterns occur with motile species (Dinophyceae, Cryptophyceae, etc.), and with non-motile forms whose physiological adaptations allow growth to large sub-surface peaks at preferred depths. These sub-surface maxima can be upwelled to the water surface, in a manner controlled by thermal stability and vertical shear, and horizontally transported to give large variations in horizontal distribution.At all latitudes diel stability cycles in surface waters can affect physiological properties important for growth, and in some circumstances can dominate the phytoplankton dynamics and distribution.Such short-term stability events merge with longer-term (e.g. annual) events with no conceptual distinction. A modern way to integrate this continuity is by scaling using spectral analysis of cyclicity. This allows biological variables (algal biomass, numbers, production) to be stochastically related to indices of stability (e.g. Brunt-Väisälä frequency).     

夏季南黄海浮游动物的垂直分布与昼夜垂直移动

对 2 0 0 1年 8月南黄海青岛外海至济州岛断面 ,浮游动物的日垂直变化、断面垂直分布及其与海区水文特征的关系进行了研究。共鉴定浮游动物 40种。断面浮游动物的平均丰度为 13 62 ind/ m 3 ;种类丰富度与暖水种数呈正相关 ,分布趋势为测区东部高、西部低 ;优势种为中华哲水蚤 (Calanus sinicus)、小拟哲水蚤 (Paracalanus parvus)、拟长腹剑水蚤 (Oithona similis)、强壮箭虫 (Sagitta crassa)、近缘大眼剑水蚤 (Corycaeus affinis)。在不同水层 ,浮游动物丰度分布不同 ,随水深增加而减少 ,表层丰度最高 (3 2 2 1ind/ m3) ,温跃层和底层分别为 743 ind/ m3、43 8ind/ m3;种类组成也不相同 ,表层数量最多的种类为鸟喙尖头(Penilia avirostris)、温跃层及其以下水体为中华哲水蚤 ,其它主要种类亦表现不同水层取向。在连续观测站 ,根据浮游动物主要种类的昼夜垂直移动与温跃层关系 ,将之分为表层分布 ,近温跃层分布 ,底层分布及全水层分布 4种类型     

Variability and possible adaptive significance of day-time vertical distribution ofLeptodora kindtii (Focke) (Cladocera) in a shallow eutrophic lake

The day-time vertical distribution of different size classes ofLeptodora kindtii in Tjeukemeer was studied during two contrasting years: 1978 and 1979. During the first year (1978) the predation pressure exerted by the planktivorous fish was rather high, whilst this was ca. 15 times lower during 1979. The diel vertical migration of different size classes ofLeptodora, was studied twice during 1979. Both studies support the predator avoidance hypothesis and also emphasize the idea that diel vertical migration is a dynamic rather than a fixed behaviour trait within a population.     

Population ecology, vertical migration and feeding of the Ponto-Caspian invader Hemimysis anomala in a gravel-pit lake connected to the River Rhine

1. The Ponto‐Caspian invader, Hemimysis anomala, was recently found in large numbers in a gravel‐pit lake connected to the Lower Rhine. Mysids were sampled with Perspex traps between September 2002 and April 2003 to study the population dynamics, vertical migration and feeding. 2. The abundance (as catch per unit effort, CPUE) of H. anomala declined from 270 individuals (ind.) trap^?1 (4 h)^?1 in December to below 4 ind. trap^?1 (4 h)^?1 in April. Average lengths ranged from 4.9 mm in autumn to 9.9 mm in March. The length–weight relationship of virgin females changed throughout the sampling period, from a size‐corrected wet weight of 5.1 mg in September to 16.6 mg in April for a female of 7.8 mm. 3. Successive monthly samples taken over 24 h revealed that H. anomala preferred the surface at twilight and night. During dawn the mysids migrated to the middle and bottom layers and were hardly found during day. Diel vertical migration clearly depends on the proximate factor light. 4. Stomach analyses revealed that larger H. anomala preferred zooplankton, whereas small individuals fed more on phytoplankton. The seasonal comparison showed an increasing percentage of zooplankton with increasing length. The proportion of zooplankton in the stomachs of large H. anomala individuals was highest during night and lowest during day. 5. The results suggest that H. anomala may become an important link between primary/secondary production and higher trophic levels in the food web of its new environments at the Lower Rhine.     

Vertical distribution of planktonic rotifers in a karstic meromictic lake

Vertical distribution of planktonic rotifers is described in relation to temperature and oxygen in Lake La Cruz, a single-doline, closed karstic lake (121 m diameter and 25 m maximum depth) which shows iron meromixis. Samples were taken by peristaltic pumping at 10 cm depth intervals in the oxycline zone from June 1987 to September 1988. A model of rotifer vertical structure in stratified lakes is proposed. Rotifers concentrate their populations at the depths with intense gradients. As stratification develops some rotifer populations show a downward migration following the thermocline and some others show an upward migration following the oxycline. The production-respiration balance in the lake, and so the position of the oxycline with respect to the thermocline and the layer of maximum production, depends on meteorological conditions. A shift in the dominance of congeneric or related species can occur in consecutive years. In Lake La Cruz, mixing conditions and subterranean inflow in spring were much more intense in 1988 than 1987, and the distance between production and decomposition depths was smaller in 1988. Anuraeopsis miraclei, an oxycline-bound species with high abundance in 1987, was displaced by A. fissa in 1988. A. fissa, which was a metalimnetic species during early summer, reached peak densities (3 × 10⁴ ind l^–1) at the oxycline, equaling the abundance of A. miraclei the preceeding year.     

Where to stay by night and day: Size-specific and seasonal differences in horizontal and vertical distribution of Chaoborus flavicans larvae

1. Data on the distributions of pelagic and benthic Chaoborus flavicans larvae were gathered in 1994 and tested for their agreement with the predator avoidance hypotheses. The development of all Chaoborus life stages, as well as the horizontal and vertical distribution in the four larval instars, was followed from May until October. We expected the largest larvae to dwell deeper by day, thus avoiding predation by visually foraging fish.
2. In agreement with this prediction body size increased with daytime depth, and this was true both between and within instars. The migration amplitude consequently increased with larval instar.
3. There was also evidence for horizontal migration, mainly in the third but also in the fourth instar.
4. Along a horizontal transect with increasing depth, locations with many benthic larvae had fewer pelagic larvae. Oxygen concentration was a good predictor of maximum benthic larval depth for most of the season but failed to predict their distribution in autumn.     

The effects of diel changes in circulation and mixing on the longitudinal distribution of phytoplankton in a canyon‐shaped Mediterranean reservoir

1. The near‐surface distribution of phytoplankton cells along the thalweg of a canyon‐shaped reservoir (El Gergal, southern Spain) during two surveys is described and interpreted as the result of time‐varying large‐scale circulation patterns, vertical mixing processes and the physiological capacity of algal cells to regulate its position in the water column. 2. Vertical gradients of chlorophyll‐a concentration developed in the water column during the day but disappeared at night, as a result of the shoaling and deepening of the diurnal mixed layer (dml). The changes in the depth of the dml are largely controlled in El Gergal by convectively driven mixing processes. The longitudinal circulation changes, in turn, as a result of weak and diurnal land‐sea breezes. The distribution of algal cells was patchy at all times but did not change during any of the surveys. 3. An expression is proposed to estimate time scales for the development of horizontal patchiness T_P based on simple concepts of transport. It is shown that T_P is in the order of a week, indicating that horizontal patchiness does not respond immediately to hourly changes in the controlling factors. The magnitude of T_P, though, depends on how the vertical distribution of chlorophyll‐a and longitudinal currents change on subdiurnal time scales. In particular, T_P is sensitive to the lag existing between the momentum and heat fluxes through the free surface, driving circulation and vertical mixing.     

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.