A comprehensive hypothesis on the migration of European glass eels (Anguilla anguilla)

The European eel (Anguilla anguilla) is a catadromous fish that spawns in the Sargasso Sea. As larvae, eels cross the Atlantic Ocean and reach the continental slope of Europe, where they metamorphose into post‐larval glass eels. These reach the continent, where some enter fresh water, some remain in marine waters, and others move between fresh and marine waters. After 5–25 years, as adult silver eels, they migrate back from fresh water to the Sargasso Sea to spawn and die. The glass eel stage is a critical step during which the eels cross the continental shelf and recruit to estuaries, where they facultatively transition to fresh water. Extensive research has been conducted to understand the behavioural mechanisms and environmental cues that aid and guide glass eels' migration. Glass eels follow odours and salinity gradients, they avoid light, and they change orientation and depth according to the tides. Recent work revealed that European glass eels also use Earth's magnetic field and lunar cues to orient. However, while we understand many aspects of their orientation behaviour, a unifying theory describing how glass eels migrate from the continental slope to fresh water is lacking. The goal of this review is to develop a comprehensive hypothesis on the migration of European glass eels, integrating previous knowledge on their orientation behaviour with recent findings on magnetic and celestial orientation. This review follows the journey of a hypothetical glass eel, describing the nature and the role of orientation cues involved at each step. I propose that, although glass eels have the sensory capacity to use multiple cues at any given time, their migration is based on a hierarchical succession of orientation mechanisms dictated by the physical properties of the environments that they occupy: (i) lunar and magnetic cues in pelagic water; (ii) chemical and magnetic cues in coastal areas; and (iii) odours, salinity, water current and magnetic cues in estuaries.     

Inshore migration and otolith microstructure/microchemistry of anguillid glass eels recruited to Iceland

The timing of catches of anguillid glass eels and their otolith microstructure and microchemistry were studied in southwest Iceland, where the European eel, Anguilla anguilla and American eel, A. rostrata have been thought to live sympatrically, to learn about their early life history and the possible mechanism of the separation between these two species ranges. Catches at the site studied suggest that glass eels may have started upstream migration as the river temperature warmed in late June and early July. The glass eels were mitochondrially identified into two species, A. anguilla and A. rostrata, although the latter were likely hybrids between the two species based on a different study. Otolith analyses showed no sharp increases in otolith increment width or sharp decrease of otolith Sr:Ca ratio in either species, which are the characteristic changes corresponding to the onset of metamorphosis in many anguillid species including A. rostrata collected in North America and A. anguilla in Europe. The mean age at recruitment determined for the glass eels in Iceland were similar between the two species (336.6 ± 41.7 and 319.3 ± 36.0 days for A. anguilla and A. rostrata, respectively), as were their total lengths (range 58.0–78.5 mm and 58.5–73.0 mm). In addition, mean age at metamorphosis (278.0 ± 36.8 and 254.0 ± 47.7 days) and total age (372.3 ± 50.8 and 352.9 ± 42.6 days) were also similar between the two species. However, these ages of A. rostrata in Iceland were older than those in North America, and those of A. anguilla collected in Iceland were roughly intermediate between the rest of Europe and North Africa. These findings support the hypothesis that the timing of metamorphosis is a key factor for determining the place of recruitment of glass eels and maintaining the geographic separation between the two species.     

Oceanic migration and spawning of anguillid eels

Many aspects of the life histories of anguillid eels have been revealed in recent decades, but the spawning migrations of their silver eels in the open ocean still remains poorly understood. This paper overviews what is known about the migration and spawning of anguillid species in the ocean. The factors that determine exactly when anguillid eels will begin their migrations are not known, although environmental influences such as lunar cycle, rainfall and river discharge seem to affect their patterns of movement as they migrate towards the ocean. Once in the ocean on their way to the spawning area, silver eels probably migrate in the upper few hundred metres, while reproductive maturation continues. Although involvement of a magnetic sense or olfactory cues seems probable, how they navigate or what routes they take are still a matter of speculation. There are few landmarks in the open ocean to define their spawning areas, other than oceanographic or geological features such as oceanic fronts or seamounts in some cases. Spawning of silver eels in the ocean has never been observed, but artificially matured eels of several species have exhibited similar spawning behaviours in the laboratory. Recent collections of mature adults and newly spawned preleptocephali in the spawning area of the Japanese eel Anguilla japonica have shown that spawning occurs during new moon periods in the North Equatorial Current region near the West Mariana Ridge. These data, however, show that the latitude of the spawning events can change among months and years depending on oceanographic conditions. Changes in spawning location of this and other anguillid species may affect their larval transport and survival, and appear to have the potential to influence recruitment success. A greater understanding of the spawning migration and the choice of spawning locations by silver eels is needed to help conserve declining anguillid species.     

Behavioural responses of glass eels of Anguilla anguilla to non-protein amino acids

Glass eels reacted chemotactically towards five non-protein constitutional amino acids, D-glutamine, D-asparagine, D-glutamic acid, D-alanine and β -alanine, dissolved in fresh water or salt water, with behavioural thresholds ranging from 10^-9 M for the most effective (D- and β ;-alanine) to 10^-7 M for the other three. With the exception of D-asparagine in fresh water and D-alanine at concentrations ≥10^-7 M, these amino acids were strong attractants. The results are compared with previous findings on the respective L-isomers. The non-protein amino acids may influence behaviour in the search for food and the recognition of conspecifics.     

A novel artificial habitat collection device for studying resettlement patterns in anguillid glass eels

The number of glass eels Anguilla australis and A. reinhardtii caught in artificial habitat collectors, made from a PVC base and polyethylene split rope fibres, was related to the number of rope fibre tufts attached to each collector rather than collector area directly. Ageing of collectors in situ to promote algal growth enhanced the catch of glass eels. Glass eels entered the collectors at night primarily during the flood tide, and did not move into the collectors during daylight hours. Glass eel abundance increased with increasing distance from the freshwater drain located in the causeway. The artificial habitat collectors are effective for assessing relative numbers of resettling glass eels and may be useful for studying recruitment and settlement patterns of other anguillid eel species, as well as identifying areas and habitats within a catchment that provide important shelter for glass eels. Sampling glass eels can be carried out with maximum effect and minimum effort using compact, aged artificial habitat collectors on the night time flood tide when low tide coincides with dusk.     

The effect of salinity on the chemotaxis of glass eels, Anguilla anguilla, to organic earthy and green odorants

Synopsis The behaviour of upstream migrating glass eels to salt and brackish water solutions of 8 pure organic earthy and green odorants was investigated. In 35 salt water, IPMCET and D-MF are ineffective, while MMP, IBMP, MT, TMCE, ETMCE and L-MF are strongly repellent. Dissolved in brackish water, MMP and ETMCE become attractive. The strong attraction which glass eels show to earthy and green odorants emerges as the level of salinity is reduced, suggesting that these chemicals could be orienting cues in the last phase of glass eel migration.     

长江口日本鳗鲡幼体色素发育时相及其体型变化

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Osmoregulation and role of nitrate during regrowth after cutting of ryegrass (Lolium perenne)

The osmotic role of nitrate during aftermath growth of Lolium perenne L. cv. Réveille was investigated. Plants were grown from seed in a controlled environment using a liquid medium with 1.0 m M NH₄NO₃ as nitrogen source.
Eight-week-old plants were cut 4.0 cm above the root system and then harvested over a 14-day period of regrowth on the same initial nutrient solution, except that nitrate was ¹⁵N labelled. Throughout the experimental period, nitrate storage and reduction in roots were low. In stubble and especially in leaves, nitrate accumulated during the first 6 days of regrowth whereas nitrate reduction mainly occurred after this period. Analyses of carbohydrate, chloride and potassium contents in stubble and leaves showed that the accumulation of nitrate osmotically compensated for the decrease in soluble sugars during the first 6 days of regrowth.
The cumulative osmotic potential of sugars, chloride and nitrate in differently treated plants was studied in stubble and leaves. Compared with uncut plants, the lower carbohydrate concentrations found in cut plants regrowing on 1.0 m M NH₄NO₃ were compensated for by an accumulation of nitrate. During aftermath growth on low nitrogen nutrition (0.2 m M NH₄NO₃), chloride replaced nitrate, supporting the proposed osmotic function of nitrate.
It is concluded that nitrate is involved in the osmotic adjustment of ryegrass during regrowth after cutting.     

越冬过程中花椒抗寒性与组织水和渗透调节物质的变化

花椒越冬过程中的抗寒性经历了弱抗寒性时期（11月初以前的时期）、抗寒性增强期（11月初至翌年1月初）、抗寒性最强期（翌年1月份）和抗寒性减弱期（翌年2月初以后）等4个时期。花椒的束缚水与自由水比值、可溶性糖和可溶性蛋白的含量与抗寒性显著相关,而脯胺酸含量与抗寒性相关性不显著。花椒可通过增加束缚水与自由水之间比值、可溶性糖和可溶性蛋白的含量来提高抗寒性,以适应低温环境。     

Annual variability in upstream migration of glass eels in a southern USA coastal watershed

We investigated the environmental factors that affected temporal variability of eel recruitment and upstream migration in a freshwater coastal river along the southeastern US. Glass eels Anguilla rostrata were collected through ichthyoplankton sampling in the lower Roanoke River, North Carolina. Monthly samples were taken from fixed stations from May 2001 through June 2003. There was no evidence of consistent seasonal migration patterns for glass eels in Roanoke River. From May through December in 2001, glass eels were captured only during August. In 2002, glass eels arrived in February and remained in ichthyoplankton samples through October, with the exception of samples from September. Peak catch occurred in March at 4.02 ± 1.2 and declined through June to 0.18 ± 0.07 (#/1,000 m³). By August, the mean density increased to 0.96 ± 0.82 and to 3.59 ± 2.77 by October. In 2003 from January through June, glass eels were captured only during February and March. Glass eels were routinely collected when river discharge rates were <150 m³ s⁻¹. River discharge rates >650 m⁻³ s⁻¹ resulted in no glass eels in our samples. Upstream migration during 2002 was not correlated with water temperature or related to lunar phase. Glass eel freshwater upstream migration was initiated when water temperatures exceeded a threshold range of 10°C to 15°C; however, glass eels continued to migrate when water temperatures approached 30°C. The overall negative effect of river discharge suggests that changes in the water release schedules of upstream hydroelectric facilities during glass eel migration could strongly influence their recruitment success.     

不同核桃品种的耐寒性及其渗透调节机制

以‘香玲’、‘鲁W06-1’、‘西洛3号’3个核桃品种（优系）为材料,对低温胁迫各材料的耐受情况及相应渗透调节物质含量和总蛋白质表达的变化情况进行分析,以揭示低温胁迫下不同核桃品种（优系）的渗透调节机制和蛋白质组分差异。结果显示:（1）3个核桃品种（优系）休眠枝条解剖结构分析表明,‘鲁W06-1’一年生休眠枝条的木质部比率最大,‘西洛3号’的韧皮部厚度显著大于其他两个品种（优系）。（2）随胁迫温度降低和胁迫时间的延长,枝条相对电导率（REC）逐渐升高,且‘西洛3号’的相对电导率均大于‘鲁W06-1’和‘香玲’。（3）核桃枝条可溶性糖、可溶性蛋白和游离脯氨酸含量均随温度降低先增加后减小,但各品种(优系)增至最大值的温度不同,其中‘鲁W06-1’枝条积累可溶性蛋白的速度和幅度较大,且各低温胁迫处理后的枝条可溶性糖和游离脯氨酸的含量也更高;3种渗透调节物质含量之间均呈极显著正相关关系,可溶性糖与游离脯氨酸含量的相关系数达0.844,表明二者对低温胁迫的应答反应密切相关。（4）在不同胁迫温度和胁迫时间处理后,核桃枝条的蛋白质表达谱带总体相似,但检测出6条表达量明显增加的差异条带,它们的分子量范围为38.9～87.9 kD。研究表明,3个核桃品种（优系）的耐寒性强弱为‘鲁W06-1’>‘香玲’>‘西洛3号’;低温胁迫下,核桃枝条迅速积累可溶性蛋白,然后积累可溶性糖和游离脯氨酸,并以耐寒性较强品种（优系）积累速度更快、积累量更大,且耐寒性较强的品种（优系）枝条蛋白谱带出现较多表达量增加的条带。     

Oceanic fronts in the Sargasso Sea control the early life and drift of Atlantic eels

Anguillid freshwater eels show remarkable life histories. In the Atlantic, the European eel (Anguilla anguilla) and American eel (Anguilla rostrata) undertake extensive migrations to spawn in the oceanic Sargasso Sea, and subsequently the offspring drift to foraging areas in Europe and North America, first as leaf-like leptocephali larvae that later metamorphose into glass eels. Since recruitment of European and American glass eels has declined drastically during past decades, there is a strong demand for further understanding of the early, oceanic phase of their life cycle. Consequently, during a field expedition to the eel spawning sites in the Sargasso Sea, we carried out a wide range of dedicated bio-physical studies across areas of eel larval distribution. Our findings suggest a key role of oceanic frontal processes, retaining eel larvae within a zone of enhanced feeding conditions and steering their drift. The majority of the more westerly distributed American eel larvae are likely to follow a westerly/northerly drift route entrained in the Antilles/Florida Currents. European eel larvae are generally believed to initially follow the same route, but their more easterly distribution close to the eastward flowing Subtropical Counter Current indicates that these larvae could follow a shorter, eastward route towards the Azores and Europe. The findings emphasize the significance of oceanic physical–biological linkages in the life-cycle completion of Atlantic eels.     

Water temperature and upstream migration of glass eels in New Zealand: implications of climate change

Glass eels migrating upstream in a New Zealand river showed a clear preference for water temperatures between 12 and 20°C, with an optimum of 16.5°C. Water temperatures <12°C and >22°C almost completely inhibited migration, which implies that warmer temperatures associated with global climate change might have a detrimental impact on glass eel recruitment in their current ranges. We established this by trapping glass eels of shortfin, Anguilla australis, and longfin, A. dieffenbachii, eels nightly from September to November. Eels caught in 2001 (50,287) outnumbered those caught in 2002 (19,954); shortfin glass eels dominated catches in both years, comprising 91–93% of the catch. Longfins were larger than shortfins, and size and pigmentation in both species increased as the seasons progressed. Temperatures within the migratory season in 2001 showed ∼14-day intervals between maxima that appeared to be associated with the new and full moons.     

Thermal preferenda and behavior of Atlantic eels (genus Anguilla) in relation to their spawning migration

Synopsis The final preferred temperatures (FPTs) of adult premigratory and migratory life-history phases of American eels, Anguilla rostrata, were determined by chronic tests in a horizontal thermal gradient. Mean FPTs were between 17 and 20°C and were not significantly different between life-history phases, acclimation temperatures, illumination regimes, photoperiods or sexual maturation states. Thermal behavior of eels was highly variable, both among individuals of the various test groups and among repeated tests of single individuals. Light inhibited behavioral thermoregulation by promoting shelter-seeking. The following inferences are drawn from the laboratory findings and observations of migrating A. rostrata and A. anguilla (European eels) in the North Atlantic: (1) decreasing temperatures may initiate downstream migration of silver eels, (2) eels may select temperatures close to their FPT in thermally stratified environments, but will tolerate higher and lower temperatures depending on illumination or other physical constraints, (3) the oceanic phase of the migration to the Sargasso Sea may take place at relatively shallow depths in the open ocean, probably within the upper 1000 meters. The strong eurythermality observed in eels may facilitate their occupation of and migration through thermally diverse and unpredictable habitats.     

Purification and further characterization of a glycoprotein from the skin mucus of Japanese eels

A glycoprotein (EGP) was purified from the skin mucus of Japanese eel Anguilla japonica . Apparent average molecular mass of the EGP was estimated to be 500 000. The EGP was found to contain 30·8% NeuAc, 26·4% GalNAc, 6·4% Gal, 0·4% NeuGc and 25·1% Thr‐rich protein. EGP was treated with alkaline borohydride for the release of carbohydrate chains (oligosaccharide alditols). Three carbohydrate chains were isolated from the released carbohydrate chains by Sephadex G‐25 (superfine) gel filtration and HPLC. Using ¹H‐NMR spectroscopy, methylation analysis and glycosidase digestion, the structures of the three carbohydrate chains were determined to be NeuAcα2→6GalNAc‐ ol , NeuAcα2→3GalNAc‐ ol and NeuAcα2→6(GalNAcα1→3)GalNAc‐ ol . An overall structure for the sialoglycoprotein from the skin mucus is proposed: the molecule is considered to consist of highly glycosylated 10 glycopeptide units (containing >40 carbohydrate chains) that are linked to the hydroxyl amino acids and spaced on average two amino acids apart.     

Seasonal changes in ionoregulatory variables of the glass eel Anguilla anguilla following estuarine entry: comparison with resident elvers

In the present study, glass eels Anguilla anguilla in the Minho River estuary (41·5° N, 8·5° W) decreased in size (standard length, L _S and mass, M ) from the beginning (autumn) to the end of the sampling season (summer). On the other hand elvers increased in L _S and M from spring to summer and were significantly larger than glass eels in paired comparisons. Branchial Na⁺/K⁺-ATPase and vacuolar (V-type) proton ATPase ( in vitro activities), two important ion transporting pumps, did not show significant seasonal changes in either glass eels or elvers although in glass eels Na⁺/K⁺-ATPase (activity) expression was significantly higher than in elvers. In a single month comparison Na⁺/K⁺-ATPase branchial mRNA expression was also higher in glass eels as was the protein level expression of both Na⁺/K⁺-ATPase and NKCC (Na⁺:K⁺:2Cl⁻ co-transporter). Immunofluorescence microscopy indicated apical CFTR Cl⁻ channel labelling in Na⁺/K⁺-ATPase positive chloride cell in glass eels which was absent in elvers. Whole body sodium concentration and percentage water did not show significant seasonal differences in either glass eels or elvers although there were significant differences between these two groups during some months.     

Re-examination of the spermatozol ultrastructure of eels: observations of the external morphology of spermatozoa in three species

The nuleus of the spermatozoa in both Anguilla anguilla and A. japonica species is asymmetric on the longitudinal axis, being gently curved and elongated with a hook-shaped superior end directed inside the crescent. A single spherical mitochondrion is located at the superoanterior portion of the nucleus. Nine striae extending from the basal portion of the flagellum run along the right side of nuclear surface. These are always oriented toward the mitochondrial portion and their ends are located on the mitochondrion. These features are seen not only in both Anguilla species but also in Muraenesox cinereus and in other species including other genera of Anguilliformes. It is suggested that these features be considered as common to Anguilliformes.     

Osmoregulation in two aquatic oligochaetes from habitats with different salinity and comparison to other annelids

The osmoregulatory capacity of two oligochaete species, Enchytraeus albidus Henle, 1837, and Heterochaeta costata (Claparède, 1863), was investigated by direct measurements of the osmolality of the coelomic fluid. Terrestrial and marine (28 S) populations of Enchytraeus albidus and a brackish water population (14 S) of H. costata were used in the study. The range of salinity acclimation investigated was 0–40. The response to osmotic stress was measured (a) after a long-term maintenance (>14 days) in various salinities (E. albidus only), and (b) after a hyperosmotic shock as a short-term time-course sequence. The rate of water loss following a hyperosmotic shock was measured for E. albidus. Long-term acclimation. E. albidus maintained a hyperosmotic coelomic fluid over all salinities tested. In low salinities the osmolality of the coelomic fluid of the marine population was significantly higher than that of the terrestrial population. Possible genetic discrepancies or long-term acclimation may account for this difference. The coelomic fluid of H. costata was hyperosmotic at 15%. S and isoosmotic at 30 S. Short-term acclimation (hyperosmotic shock). Both species investigated, kept at 15 S and then exposed to a salinity of 30, showed fast responses: within the first two hours the internal concentrations were adjusted to the new external condition with only small subsequent changes. Regulation of the body-water content after an exposure to a hyperosmotic shock was much slower: individuals of terrestrial E. albidus, acclimated for two weeks to either 0 or 15 S, had the same water content; hence, they showed a 100% regulation. However, after exposure to 30 S, a 100% regulation was still not attained 4 days after the hyperosmotic shock. Enchytraeus albidus is capable of actively reducing water loss following the hyperosmotic shock: the observed loss of water was only 40% of that expected for a passive osmotic flow. The observed reactions are compared with those in other annelids. It seems that an active transport of ions combined with a changeable permeability of the body wall play a major role in the regulation of body fluids.     

Age and growth of eels Anguilla japonica in a Chinese river

Age and growth of Anguilla japonica sampled from the Qiulu River in southern China were studied. Mean body lengths and weights of the eels at each age were back-calculated from their otoliths. The growth parameters for the von Bertalanffy function gave L _∞= 98–2cm, K =0·07 and t ₀=− 1·94. Scales were observed to form firstly at the middle part of the lateral line, at a minimum body length of 160 mm and were inappropriate for ageing.     

Osmoregulation by the aphid Myzus persicae: A physiological role for honeydew oligosaccharides

The dry matter content of honeydew produced by Myzus persicae feeding on artificial diets increased with increasing sucrose concentrations of the diet. Whereas the diet osmolalities ranged from 828 to 1800 milliosmolal, the honeydew osmolality was relatively constant (about 500 milliosmolal). This osmoregulatory capacity is achieved largely by variations in the mean molecular weight of glucose-containing honeydew oligosaccharides.