The ontogeny of schooling behaviour in the striped jack

The ontogeny of schooling behaviour was investigated in laboratory-reared striped jack Pseudocaranx dentex larvae and juveniles. Mean separation angle between nearest individuals averaged 78 and 82° in 10- and 12-mm fish, respectively, and decreased to 57° in 16-mm fish. In addition, interindividual distance decreased from three times body length in 12-mm fish to twice the body length in 16-mm individuals; this value continued to decrease gradually to 1·3 at 19 mm to 0·79 at 30 mm. Mutual attraction due to visual recognition was observed in 12-mm fish and increased with total length. The optokinetic response ( R_o , the ability of a fish to keep station with moving patterns) first appeared in 4–6-mm larvae. In spite of the traditional emphasis of the importance of R_o in schooling behaviour, the onset of R_o was insufficient to result in school formation. The emergence of mutual attraction seems essential for the ontogeny of schooling.     

Critical involvement of dietary docosahexaenoic acid in the ontogeny of schooling behaviour in the yellowtail

To investigate involvement of the central nervous system in the ontogeny of behaviour, diets of differing quality were used to rear yellowtail Seriola quinqueradiata larvae and juveniles. Artemia nauplii enriched with oleic acid (OA), eicosapentaenoic acid (EPA), or two different concentration levels of docosahexaenoic acid (DHA) were fed to yellowtail larvae (L^T 7 mm; 13 days old) for 12 days, and their behavioural development was analysed together with growth, survival, activity and fatty acid composition. Yellowtail fed with DHA–enriched Artemia showed mutual attraction behaviour at 11 mm L^T, while those fed with OA– or EPA–enriched Artemia did not show this behaviour at the same size. While fish in the OA group showed poor growth, survival and activity index, fish in the EPA group were similar to the two DHA groups. In addition, most fish tested, including the OA group, showed clear optokinetic responses. Fatty acid composition of the diet was reflected in that of the fish body. Therefore, dietary DHA in the larval stage is considered to be essential for the development of schooling behaviour in the yellowtail.     

大麻哈鱼卵黄囊期仔鱼异速生长及其生态学意义

运用实验生态学的方法, 对大麻哈鱼(Oncorhynchus keta Walbaum)卵黄囊期仔鱼的异速生长及器官优先发育在早期生存和环境适应上的生态学意义进行了研究。结果表明, 大麻哈鱼卵黄囊期仔鱼的感觉、摄食, 呼吸和游泳等器官快速分化, 许多关键器官均存在异速生长现象。在身体各部分中, 头部和尾部为正异速生长, 躯干部为负异速生长, 体高有先增大后减小的趋势; 在头部器官中, 眼径、口宽、吻长和眼后头长均为正异速生长; 在游泳器官中, 胸鳍、腹鳍、背鳍、臀鳍、背鳍基、臀鳍基和尾鳍均为正异速生长, 脂鳍为负异速生长, 其中, 腹鳍在全长25.31 mm、12日龄出现生长拐点, 但拐点前后均为正异速生长。大麻哈鱼卵黄囊期仔鱼感觉、摄食, 呼吸和游泳等器官的快速发育, 使出膜后的仔鱼在最短的时间内获得了与早期生存密切相关的各种能力, 对适应复杂多变的外界环境具有重要的生态学意义。       

Ontogeny of swimming speed, schooling behaviour and jellyfish avoidance by Japanese anchovy Engraulis japonicus

The ontogeny of swimming speed, schooling behaviour and jellyfish avoidance was studied in hatchery-reared Japanese anchovy Engraulis japonicus to compare its life-history strategy with two other common pelagic fishes, jack mackerel Trachurus japonicus and chub mackerel Scomber japonicus. Cruise swimming speed of E. japonicus increased allometrically from 1·4 to 3·9 standard length (L(S) ) per s (L(S) s(-1) ) from early larval to metamorphosing stage. Burst swimming speed also increased from 6·1 to 28 L(S) s(-1) in these stages. Cruise speed was inferior to that of S. japonicus, as was burst speed to that of T. japonicus. Engraulis japonicus larvae were highly vulnerable to predation by moon jellyfish Aurelia aurita and were readily eaten until they reached 23 mm L(S) , but not at 26 mm L(S) . Schooling behaviour (indicated by parallel swimming) started at c. 17 mm L(S) . Average distance to the nearest neighbour was shorter than values reported in other pelagic fishes. The relatively low predator avoidance capability of E. japonicus may be compensated for by their transparent and thus less conspicuous body, in addition to their early maturation and high fecundity.     

Morphogenesis in hatchery-reared larvae of the black rockfish,Sebastes schlegeli,and its relationship to the development of swimming and feeding functions

The developmental sequence of morphological characteristics related to swimming and feeding functions was investigated in hatchery-reared larvae and juveniles ofSebastes schlegeli, a viviparous scorpaenid. The fish were extruded at an early larval stage, when the mean body size was 6.23 mm TL. Fin-ray rudiments became visible at 9.0 mm TL in the dorsal and anal fins, at 8.0 mm TL in the pectoral and pelvic fins and 6.0 mm TL (size at extrusion) in the caudal fin. Completion of segmentation of soft rays in the dorsal and anal fins was attained by 14 mm TL and in all fins by 17 mm TL. Branching of soft rays in the respective fins started and was completed considerably later than the completion of segmentation, as well as ossification of the fin-supports. Morphological transformation from larva to juvenile was apparently completed by about 17 mm TL. Although the completion of basic juvenile structures was attained by transformation at that body size, succeeding morphological changes occurred between 17 mm and 32 mm TL. Newly-extruded larvae possessed one or two teeth on the lower pharyngeal and pharyngobranchials 3 and 4, but lacked premaxillary, dentary, palatine and prevomer teeth. The fish attained full development of gill rakers and gill teeth by 15 mm TL, the upper and lower pharyngeal teeth subsequently developing into a toothplate. Development of the premaxillary, dentary and palatine teeth was completed at about 30 mm TL, by which time loop formation of the digestive canal and the number of pyloric caeca had attained the adult condition. The developmental sequence of swimming and feeding functions during larval and early juvenile periods appeared to proceed from primitive functions to advanced or complex ones, from the ability to produce propulsive force to that of swimming with high maneuverability and from development of the irreducible minimum function of passing food into the stomach to the ability to actively capture prey via passive food acquisition with the gill rakers and gill teeth. The relationship of morphological development to the behavior and feeding activity of artificially-produced hatchlings is also discussed.     

斑鰶仔、稚鱼形态发育及异速生长模式研究

运用光学显微镜技术和实验生态学方法, 对斑鰶(Konosirus punctatus)早期形态发育观察、异速生长模式及其生态学意义进行了研究。结果表明: 在水温(21.5±0.5)℃下, 初孵仔鱼全长(3.18±0.52) mm, 斑鰶仔鱼期从孵化出膜到43日龄棱鳞开始出现前, 稚鱼期从44日龄棱鳞出现到55日龄全身覆满鳞片。斑鰶的形态变化和器官分化主要发生在仔鱼期。斑鰶的吻长、躯干长、肠道长、胸鳍长、腹鳍长等重要形态学指标均存在异速生长现象, 其生长拐点依次为 42日龄(TL: 26.41 mm)、24日龄(TL: 15.57 mm)、31日龄(TL: 21.41 mm)、41日龄(TL: 25.47 mm)、42日龄(TL: 26.41 mm)。相对于全长、吻长和胸鳍长在拐点前后由正异速生长变为等速生长, 腹鳍长由正异速生长转为负异速生长, 这为呼吸、摄食和成功逃避捕食者提供有利条件; 而肠道长由负异速生长变为等速生长, 这可能与斑鰶的食性转化有关。综上所述, 为适应复杂多变的生存环境, 斑鰶在早期发育阶段优先发育对生长生存起关键作用的器官, 这对提高仔、稚鱼的存活率具有重要的生态学意义。研究将为今后进一步人工繁育和苗种培育提供理论依据。     

眼斑双锯鱼仔稚鱼发育异速生长

运用生态学和传统理论生物学的研究方法, 对孵化后眼斑双锯(Amphiprion ocellaris)仔、稚鱼在早期生存和环境适应上的异速生长及器官优先发育生态学意义进行了研究, 以期为眼斑双锯鱼人工繁殖和育苗提供参考资料。以11日龄为眼斑双锯鱼仔、稚鱼的区分时期, 结果表明, 眼斑双锯鱼仔、稚鱼的感觉、摄食和游泳等器官快速分化, 均存在异速生长现象。在头部器官中, 吻长、眼间距、口宽和头高在仔鱼期均为正异速生长, 吻至鳃裂前缘长和眼径为负异速生长。在身体各部位中, 仔鱼期体高、躯干长、尾长、尾柄长、尾柄高和体厚均为正异速生长, 仅头长为负异速生长; 在游泳器官中, 仔鱼期眼斑双锯鱼尾鳍、背鳍、胸鳍、腹鳍和臀鳍均为正异速生长。稚鱼期眼斑双锯鱼头部、躯干及游泳等各器官均为负异速生长。眼斑双锯鱼这些关键器官的异速发育, 对适应环境因子变化具有重要的生态学意义。     

Early developmental and allometric patterns in the electric yellow cichlid Labidochromis caeruleus

The electric yellow cichlid Labidochromis caeruleus is a mouth‐brooding haplochromine cichlid from Lake Malawi and one of the most popular cichlids in the ornamental fish industry. To investigate the early development of L. caeruleus from hatching until the juvenile stage, we studied its morphological development and allometric growth patterns. In newly‐hatched larvae, most organs and body parts were not yet differentiated and continued to develop until 15 days post hatching (dph). The yolk sac was depleted at 13 dph. There was allometric growth, primarily in the anterior and posterior regions of the body, and inflection points when trajectories of allometric growth changed. Head and tail growth was prioritized, suggesting that body parts linked to feeding and swimming behaviour mature earlier than the rest of the body. Additionally, growth patterns revealed that development of organs related to vital functions such as branchial respiration, sensation, exogenous feeding and swimming was prioritized. Comparisons with other African and Neotropical cichlids revealed differences in ontogenetic processes and allometric growth along the anterior–posterior axis as well as variation in developmental timing. These results indicate how early morphological development and ontogenic processes might respond to the distinctive parental care observed in mouth‐brooding cichlids.     

Allometric growth and larval development in Pacific red snapper Lutjanus peru under culture conditions

Allometric growth is a common feature during fish larval development. It has been proposed as a growth strategy to prioritize the development of body segments related to primordial functions like feeding and swimming to increase the probability of survival during this critical period. In the present study we evaluated the allometric growth patterns of body segments associated to swimming and feeding during the larval stages of Pacific red snapper Lutjanus peru. The larvae were kept under intensive culture conditions and sampled every day from hatching until day 33 after hatching. Each larva was classified according to its developmental stage into yolk-sac larva, preflexion larva, flexion larva or postflexion larva, measured and the allometric growth coefficient of different body segments was evaluated using the potential model. Based on the results we can infer the presence of different ontogenetic priorities during the first developmental stages associated with vital functions like swimming during the yolk-sac stage [total length (TL) interval = 2.27–3.005 mm] and feeding during the preflexion stage (TL interval = 3.007–5.60 mm) by promoting the accelerated growth of tail (post anal) and head, respectively. In the flexion stage (TL interval = 5.61–7.62 mm) a change in growth coefficients of most body segments compared to the previous stage was detected, suggesting a shift in growth priorities. Finally, in the postflexion stage (TL interval = 7.60–15.48 mm) a clear tendency to isometry in most body segments was observed, suggesting that growth priorities have been fulfilled and the larvae will initiate with the transformation into a juvenile. These results provide a framework of the larval growth of L. peru in culture conditions which can be useful for comparative studies with other species or in aquaculture to evaluate the changes in larval growth due to new conditions or feeding protocols.     

鲈鲤仔鱼的异速生长模式

采用实验生态学方法研究了鲈鲤(Percocypris pingi pingi)仔鱼(0～57日龄)的异速生长模式.结果显示:鲈鲤仔鱼全长由慢速生长到快速生长的转折点为25日龄;其多数外部器官均具有异速生长特点,头部和尾部的生长快于躯干部,均在22 ～ 27日龄出现生长拐点;眼径在14 ～ 15日龄较早出现生长拐点,促使眼睛充分发育,以提高早期仔鱼开口期摄食外源食物的能力;吻长在33～34日龄出现生长拐点,促进了口的充分发育,以适应不同的饵料环境;胸鳍、背鳍、尾鳍、臀鳍和腹鳍分别在13～14日龄、31～32日龄、32 ～33日龄、38 ～39日龄、43 ～ 44日龄出现生长拐点,除胸鳍和尾鳍外,其余各鳍的鳍条均在拐点处分化完全,即鲈鲤仔鱼的游泳能力已得到大幅提高.研究表明,鲈鲤仔鱼的异速生长模式,保证了各重要功能器官的充分发育,以适应多变的环境,有效地保障了其早期的生存,可为育苗生产和野生早期资源的保护提供技术支撑.     

The effect of dietary docosahexaenoic acid on schooling behaviour and brain development in larval yellowtail

The onset of schooling behaviour and brain growth in larval yellowtail Seriola quinqueradiata larvae fed on Artemia enriched with oleic acid (OA), eicosapentaenoic acid (EPA) or docosahexaenoic acid (DHA) for 10 days was investigated. Larvae from the DHA groups schooled with parallel orientation while those of the EPA groups showed only aggregation. Larvae of the OA groups were dispersed. DHA and EPA groups showed significantly smaller nearest neighbour distance compared with OA groups, and DHA groups showed significantly smaller values of nearest neighbour angle compared with the other groups. The relative volume of the tectum opticus (TO) of the DHA group was significantly larger in two experiments and also the relative volume of the cerebellum (CE) of that group was significantly larger in one experiment. Dietary DHA is probably critically important for the development of the brain, particularly TO, during the larval stages, and that development of TO may be the key factor of the ontogeny of schooling behaviour.     

拟赤梢鱼的胚胎发育和仔稚鱼生长特性观察

为阐明拟赤梢鱼(Pseudaspius leptocephalus)胚胎发育和仔稚鱼发育特点,采用人工催产的方式获得受精卵,观察分析了拟赤梢鱼胚胎发育和仔稚发育的时序特征.结果表明:拟赤梢鱼成熟卵粒为黄色圆球形,平均卵径为(1.77±0.20)mm,遇水具微黏性;在水温23℃条件下,胚胎发育经历合子期、卵裂期、囊胚期、...     

Development of eggs,larvae and juveniles of the labrid fish,Halichoeres poecilopterus,reared in the laboratory

Embryonic, larval and juvenile development of the labrid fish,Halichoeres poecilopterus, is described using a laboratory-reared series. The eggs, measuring 0.60–0.72 mm in diameter, were pelagic and spherical with a single oil globule (0.12–0.16 mm in diameter). Hatching occurred 18 h 48 min after spawning. The newly-hatched larvae, measuring 1.46–1.70 mm TL, had 8–114 + 16–18 myomeres. A conspicuous melanophore appeared on the dorsal finfold 8 h after hatching, at ca. 2 mm TL. The yolk was completely absorbed 3 days after hatching, at 2.52–2.72 mm TL. Flexion of the notochord started at ca. 6 mm TL and was finished at ca. 8 mm TL. Aggregate numbers of all fin rays were completed at ca. 14 mm TL. Squamation was almost completed at ca. 20 mm TL.     

Development of the escape response in teleost fishes: do ontogenetic changes enable improved performance?

Teleost fishes typically first encounter the environment as free-swimming embryos or larvae. Larvae are morphologically distinct from adults, and major anatomical structures are unformed. Thus, larvae undergo a series of dramatic morphological changes until they reach adult morphology (but are reproductively immature) and are considered juveniles. Free-swimming embryos and larvae are able to perform a C-start, an effective escape response that is used evade predators. However, escape response performance improves during early development: as young fish grow, they swim faster (length-specific maximum velocity increases) and perform the escape more rapidly (time to complete the behavior decreases). These improvements cease when fish become juveniles, although absolute swimming velocity (m s(-1)) continues to increase. We use studies of escape behavior and ontogeny in California halibut (Paralichthys californicus), rainbow trout (Oncorhynchus mykiss), and razorback suckers (Xyrauchen texanus) to test the hypothesis that specific morphological changes improve escape performance. We suggest that formation of the caudal fin improves energy transfer to the water and therefore increases thrust production and swimming velocity. In addition, changes to the axial skeleton during the larval period produce increased axial stiffness, which in turn allows the production of a more rapid and effective escape response. Because escape performance improves as adult morphology develops, fish that enter the environment in an advanced stage of development (i.e., those with direct development) should have a greater ability to evade predators than do fish that enter the environment at an early stage of development (i.e., those with indirect development).     

The onset of piscivory in a freshwater fish species: analysis of behavioural and physiological traits

The onset of piscivory in fish, resulting in a shift from zooplankton or invertebrate to fish prey, was studied on pikeperch (Sander lucioperca) larvae displaying and not displaying piscivorous behaviour at different ages (23, 30, 37, 44 and 52 days post-hatching) using behavioural (attack, capture and swimming activity), morphological (allometry) and digestive enzymatic (trypsin, α-amylase and pepsin) analyses. The shift from zooplanktonic food items (Artemia nauplii) to a piscivorous diet did not occur at the same time for all individuals within the same cohort. Predation tests, conducted under controlled conditions (20°C; ad libitum feeding), showed that some larvae attacked fish prey as early as the age of 3 weeks [11.0 ± 1.3 mm total length (TL)], whereas others did not start until the age of 6 weeks (16.6 ± 1.9 mm TL). Piscivorous individuals were bigger, with larger heads, longer tails, higher acid protease and lower alkaline protease activities, than non-piscivorous conspecifics. In conclusion, high interindividual variability in morphological and digestive system developments linked to the development of predatory abilities could induce cannibalism in fish.     

日本七鳃鳗胚胎发育及卵黄囊期仔鱼的异速生长

通过观察日本七鳃鳗Lampetra japonica (Martens, 1868)胚胎外部形态和内部组织结构变化, 描述受精卵从卵裂至器官形成以及仔鱼孵出的发育阶段, 采用实验生态学方法研究卵黄囊期仔鱼的异速生长模式。结果表明: 日本七鳃鳗卵子为乳白色, 呈卵圆形; 受精卵卵裂方式为全卵裂; 胚胎发育过程主要包括卵裂期、囊胚期、原肠胚期、神经胚期、头凸期、孵出前期和孵出期, 历时11—12d; 初孵仔鱼为乳白色, 全长约(3.41±0.24) mm, 体质量约为0.0006 g。日本七鳃鳗胚胎发育研究可为了解七鳃鳗胚胎发育过程, 早期脊椎动物的起源和发育进化研究提供参考。卵黄囊期内仔鱼身体各部分中, 头长和尾长均表现出快速生长, 同在7 日龄出现生长拐点, 且生长拐点后的生长速率都大于生长拐点前的生长速率; 而仔鱼体长在卵黄囊期内表现出慢速生长。在头部器官中, 吻长、鳃前长和鳃长均表现出快速生长现象, 吻长和鳃长分别在9日龄和8日龄出现生长拐点; 口笠长在3日龄时出现生长拐点, 在生长拐点前为等速生长, 而在生长拐点后表现出快速生长; 眼径和眼鳃间距则分别表现出等速生长和慢速生长。泄殖孔在卵黄囊期内未出现生长拐点, 生长速率相对于全长生长速率表现出快速生长现象。七鳃鳗卵黄囊期仔鱼的异速生长是在长期进化过程中, 适应早期生活环境和独特的生活方式而形成特有的发育机制。     

红鳍笛鲷仔、稚鱼异速生长

运用生态学和传统理论生物学的研究方法,对孵化后红鳍笛鲷（Lutjanus erythropterus）仔、稚鱼在早期生存和环境适应上的异速生长及器官优先发育生态学意义进行了研究,以期为红鳍笛鲷人工繁殖、育苗提供参考资料。以17日龄为红鳍笛鲷仔、稚鱼的区分时期,结果表明,红鳍笛鲷仔、稚鱼的感觉、呼吸摄食和游泳等器官快速分化,均存在异速生长现象。在头部器官中,吻长、口宽、眼径和头高在仔鱼期均为正异速生长,稚鱼期吻长为等速生长,口宽、眼径和头高为负异速生长。在身体各部位中,仔鱼期头长和体高为正异速生长,躯干部和尾长为负异速生长;稚鱼期体高和躯干长为正异速生长,头长和尾长为等速生长;在游泳器官中,仔鱼期红鳍笛鲷背鳍、腹鳍、尾鳍为正异速生长,胸鳍为等速生长,稚鱼期臀鳍为正异速生长,腹鳍、胸鳍和尾鳍为等速生长,背鳍为负异速生长。红鳍笛鲷这些关键器官的快速发育,使外源性营养开始后以最小的代谢损耗获得了生存能力的显著提升,对挑战和适应纷繁变换的外界压力具有重要的生态学意义。     

Embryonic and morphological development of larvae and juveniles of the Amberjack,Seriola dumerili

Embryonic and morphological development of larvae and juveniles of the amberjack,Seriola dumerili Risso, are described using specimens raised at Yaeyama Station (Ishigaki Island, Okinawa Pref.), Japan Sea Farming Association. The specimens obtained from brood fish (3 females, 3 males) were treated with gonadotropin and spawned on 6th of April 1987. The eggs of amberjack are pelagic, spherical in shape and 1.01–1.17 mm in diameter. The yolk is roughly segmented and has a single oil globule 0.22–0.24 mm in diameter. The perivitelline space is narrow. During development, a few melanophores and no xanthophores were observed on yolk. Hatching took place 35 hrs. 15 min. after spawning out at temperatures 23.1–23.7°C. The newly hatched larvae were 2.84–3.04mm in TL with 27 (13+14) myomeres and an oil globule anteriorly situated beyond the head. 3 days after hatching 4.00 mm TL, the mouth opened. 10 days after hatching 4.26 mm TL, small denticles appeared on the margin of the upper jaw and there were 1 anterior and 2 posterior preopecular spines. At 5.96mm TL, notochord was slightly flexed. Caudal, dorsal and anal fins with rudiments of rays appeared at 8.00 mm TL. The specific numbers of all fin rays and spines were obtained in a juvenile 9.60 mm TL. In a juvenile 34.25 mm TL, 54 days after hatching, the characteristic brown band of amberjack had appeared on head. Some notable changes in relative growth were observed at 5 mm and 15 mm in TL.     

Development of eggs, larvae and juveniles of laboratory-reared blue whiting,Sillago parvisquamis (Percoidei: Sillaginidae)

The embryonic, larval and juvenile development of blue whiting,Sillago parvisquamis Gill, are described from a series of laboratory-reared specimens. Mean egg diameter and mean total length (TL) of newly-hatched larvae were 0.71 mm and 1.58 mm, respectively. The eggs were non-adhesive, buoyant and spherical with an oil globule (mean diameter 0.18 mm). Hatching occurred about 20 hours after fertilization at a temperature of 24.0–25.0°C, newly-hatched larvae having 38–40 myomeres. The yolk and oil globule were completely absorbed 3 days after hatching at 2.8–3.2 (mean 3.0) mm TL. Notochord flexion was completed by 7.2–8.2 (7.7) mm TL, and pectoral and caudal fin rays fully developed by approximately 10 mm and 8.5 mm TL, respectively. Completion of fin development occurred in the following sequence: caudal, pectoral, anal and second dorsal, first dorsal and pelvic, the last-mentioned by approximately 11 mm TL. The larvae ofS. parvisquamis andS. japonica, which closely resemble each other in general morphology and pigmentation, could be distinguished as follows. Newly-hatchedS. parvisquamis larvae had more myomeres thanS. japonica (38–40 vs. 32–34) and more melanophores on the dorsal surface of the body (19–28 vs. about 40).Sillago japonica had a vertical band of melanophores on the caudal peduncle, which was lacking in postflexionS. parvisquamis larvae. In addition, juveniles ofS. parvisquamis (larger than 23 mm TL) had melanophores on the body extending anteriorly to below the lateral line to form a midlateral band, whereas no obvious band occurred on similarly-sizedS. japonica juveniles.     

The development of schooling behaviour in Atlantic herring Clupea harengus

The timing of the development of schooling behaviour in Atlantic herring Clupea harengus and the quantitative and qualitative evolution of such behaviour in the larval and post-metamorphic stages were investigated with a simulation model and experimental observations of laboratory-reared fish. Herring larvae started to develop schooling behaviour between the lengths of 35 and 40 mm. This coincided with the beginning of metamorphosis, characterized by ontogenetic changes which require a switch to a different antipredator strategy from that employed by early larvae, while providing the sensory and anatomical mechanisms necessary for the formation of schools. Schooling behaviour was established at the end of metamorphosis (50–55 mm) and its characteristics continued to evolve throughout the early juvenile stage. A critical period in terms of vulnerability to predation is expected between the end of the larval and beginning of the juvenile stages, before schooling behaviour is fully developed. The implications of some of the results for the methodology of future studies are also discussed.