Comparison of otolith growth and morphology with somatic growth and age in young-of-the-year bluefin tuna

Otolith morphological characteristics were studied using image analysis techniques and the relationships between otolith growth and somatic growth and age, as estimated from counting daily otolith increments, were examined in young-of-the-year (YOY) bluefin tuna Thunnus thynnus ranging in fork length ( L _F) from 8·5 to 55·5 cm. Whole otolith length, width, area and perimeter, and three shape indexes, circularity, E value and rectangularity, were extracted for each pair of sagittae. Since no statistical significant differences between left and right otolith morphometrics were found, only one otolith from each fish was used for correlations. Statistically significant relationships were observed between otoliths measurements and fish somatic growth when a linear regression was applied after logarithmic transformation of all variables tested. Among the variables, otolith length was the one that showed the highest correlation with L _F, followed by otolith area and perimeter, whereas otolith rectangularity exhibited the lowest correlation. Statistically significant relationships were also observed between the otolith variables tested and the age of the fish, which ranged from 20 to 129 days. The ages estimated using otolith mass were very close to those assessed using daily increment counts (bias ranged from 1 to 24 days). Therefore, otolith mass could represent a valuable criterion for age estimation in YOY bluefin tuna that is objective, economic and easy to perform compared to daily increment counting method.     

Relationships between otolith microstructure, otolith growth, somatic growth and ontogenetic transitions in two cohorts of windowpane

Otolith increments in larval and juvenile windowpane Scophthalmus aquosus can provide an estimate of daily age for spring-spawned individuals held under summer conditions. Otolith increments for spring- and autumn-spawned individuals occurred at intervals >1 day under winter conditions. A significant decrease in the slope of the linear relationship between otolith size and somatic size at 40 mm L_S coincided with significant habitat, morphological and behavioural transitions. In smaller, field-captured windowpane belonging to spring- and autumn-spawned cohorts, the formation of accessory growth centres coincided with a transitional settlement period and the completion of eye migration (c. 8–20 mm L_S). Back-calculated growth rate estimates for spring-spawned windowpane were significantly faster than those for autumn-spawned windowpane and these differences could produce differential rates of survival for the two cohorts during the first year of life.     

Growth estimates from otolith increment widths of juvenile chinook salmon (Oncorhynchus tshawytscha) reared in changing environments

For otolith increments to provide useful estimates of fish growth, otolith growth must covary closely with somatic growth. We reared groups of juvenile chinook salmon ( Oncorhynchus tshawytscha Walbaum) for 70 days, changing ration or temperature during a 20-day treatment period. Restricted rations halted somatic growth, however increment widths decreased gradually; somatic growth was overestimated from increment width. Otolith growth followed changes in water temperature more closely than changes in ration, supporting a hypothesized effect of metabolic rate on otolith growth. Increment growth was only loosely coupled to fish growth rate, and may also be affected by past growth histories. For juvenile fish, increment widths may not be sensitive indicators of short-term changes in growing conditions.     

The effect of temperature and somatic growth on otolith growth: the discrepancy between two clupeid species from a similar environment

Otolith growth rates of the early life stages of herring Clupea harengus ( n = 472) and smelt Osmerus eperlanus ( n = 348) collected in the Vistula Lagoon (Baltic Sea) during 1997–1999 were analysed. The larvae and early juveniles were not only collected in the same geographical area they were also of the same size (range 15–43 mm standard length, L _S), similar ages and were collected during the same seasons (May to July). Although the two clupeid species experienced very similar environmental conditions, there were significant discrepancies in the analysed relationships. The otolith growth of larval and juvenile smelt was very strongly related to somatic growth while temperature had a minor effect. In herring, the effect of somatic growth, although clearly visible and statistically highly significant, was of less importance than temperature. Furthermore, variation in the otolith size and L _S relationship was affected by temperature and somatic growth in both species, but the variance of otolith size at L _S was higher for herring than for smelt. Although growth backcalculation from otoliths can presently be recommended as an appropriate method for use with both smelt and herring (despite possibly lower precision and accuracy with the latter), other methods referring directly to short-term increment width changes ( e.g. marginal increment analysis) are recommended for smelt but not for herring.     

Validation of daily otolith increments in larval and juvenile Chinese sucker, <Emphasis Type="Italic">Myxocyprinus asiaticus</Emphasis>

Otolith development was observed and the formation of daily growth increments in otoliths of Chinese sucker, Myxocyprinus asiaticus, was validated by monitoring known-age larvae and juveniles in the laboratory from 2003 to 2005. Otolith shape changed with larval and juvenile development, and there was an exponential relationship until a body length of 16 mm or so, and a linear relationship after a body length of 16 mm between otolith size and fish size. The first increment was identified in larvae 1 day after hatching. The regressed equations between daily age (D) and increment number in otoliths (N) were N = −0.64 + 0.96D in lapillus, and N = −0.31 + 0.98D in sagitta. The slopes were not significantly different than 1.0. This demonstrated that otolith increments in this species were formed daily and can be used for daily age determination.     

Otolith formation,microstructure and daily increment validation in juvenile perch Perca fluviatilis

The otoliths of laboratory‐reared larval and juvenile perch Perca fluviatilis of known age were analysed to determine the age of otolith formation and validate the formation of daily increments. There was a linear relationship between number of increments and age in days post‐hatching, although by 82 days post‐hatching daily increment counts underestimated actual age by an average of 5 days. Otolith dimensions in relation to standard length indicated allometric growth of otoliths until completion of yolk absorption, and isometric growth thereafter, up to 82 days post‐hatching.     

Ontogenetic and environmental effects on otolith shape variability in three Mediterranean European eel (Anguilla anguilla, L.) local stocks

Otolith morphology is an efficient tool for the discrimination of fish stocks, populations and species when comparative genetic data are not available. Currently, the relationship between environmental factors and otolith shape is poorly characterized for the European eel (Anguilla anguilla), a highly migratory catadromous species constituting a single, randomly mating stock. The present study analyses the differences in otolith morphology between three Mediterranean eel local stocks from different environmental contexts (i.e. two brackish lagoons and one river). The relationship between otolith shape and otolith size was studied by means of Elliptic Fourier analysis and multivariate statistics. Otolith profile was digitally acquired and Cartesian coordinates were extracted. Partial Least Square (PLS) analysis pointed to continuous allometric growth in size and shape in otoliths from all three sites. In the three environments, shape variations occurred during growth as indicated by the presence of a significant and positive relationship between otolith size and the first PLS latent vector (i.e. which bears most of the information regarding otolith outline). Differences between smaller and larger sized otoliths were investigated using PLS Discriminant Analysis (PLSDA) and cluster analysis. Results indicate that otolith shape is highly uniform at smaller than at larger sizes. These shape differences apparently overlap the initial differentiation of the small otolith outlines acquired by eels during the growing phase as elvers in the marine environment. Data were discussed considering that the physical and chemical habitat variability in brackish lagoons and river could underlie a marked change in otolith shape during the animals' growth.     

An evaluation of the otolith characteristics of Conger conger during metamorphosis

A sample of 20 metamorphosing conger eel Conger conger leptocephali were collected from the Minho River, Portugal, in February 1999 and their sagittal otoliths were analysed by scanning electron microscopy. Four different etching agents were applied along both sagittal and frontal sections during otolith preparation to examine the microstructural growth in this species. Otolith growth increments were visible throughout the increment countable zone using all four treatments, but a permanent peripheral diffuse zone, where the daily increments were unclear, appeared on all otoliths, preventing accurate age estimation. To understand more about the nature of the diffuse zone, otoliths of 10 other metamorphosing leptocephali reared in aquaria were marked by immersion in tetracycline hydrochloride. The distance between the fluorescent marks and otolith edge, measured over a fixed period of time, was used to estimate the otolith growth rate. The application of this technique led to an anomalously high estimated otolith growth rate, probably as a result of the capture, marking and handling stress.     

Do otolith increments allow correct inferences about age and growth of coral reef fishes?

Otolith increment structure is widely used to estimate age and growth of marine fishes. Here, I test the accuracy of the long-term otolith increment analysis of the lemon damselfish Pomacentrus moluccensis to describe age and growth characteristics. I compare the number of putative annual otolith increments (as a proxy for actual age) and widths of these increments (as proxies for somatic growth) with actual tagged fish-length data, based on a 6-year dataset, the longest time course for a coral reef fish. Estimated age from otoliths corresponded closely with actual age in all cases, confirming annual increment formation. However, otolith increment widths were poor proxies for actual growth in length [linear regression r ² = 0.44–0.90, n = 6 fish] and were clearly of limited value in estimating annual growth. Up to 60 % of the annual growth variation was missed using otolith increments, suggesting the long-term back calculations of otolith growth characteristics of reef fish populations should be interpreted with caution.     

Temperature effects on otolith pattern formation in Atlantic cod Gadus morhua

The effect of food intake and temperature on otolith macrostructure and microstructure was examined experimentally in Atlantic cod Gadus morhua. Daily increment formation was validated and otolith accretion rate and optical density quantified using image analysis. Two‐week periods of starvation had no discernable effect on otolith increment width or optical density, despite having negative effects on somatic growth. In contrast, temperature had a strong positive effect on otolith accretion rate and clear effects on optical density with the otolith becoming more translucent at higher temperatures. Somatic growth, otolith accretion and otolith optical density each had a significantly different response curve to temperature. No relationship was detected between individual somatic growth rates and the accretion rate or optical properties of the otolith. The experimental manipulation of temperature‐induced otolith patterns similar to the ‘false ring’ secondary structures sometimes observed in the otoliths of wild fish. The results suggest that otolith pattern arises from a combination of temperature and seasonal effects, but not directly from individual variation in somatic growth.     

Growth of North Alboran Sea sardine larvae estimated by otolith microstructure, nucleic acids and protein content

Wet mass and DNA, RNA and protein content increased significantly with standard length ( L _S) of sardine Sardina pilchardus larvae, collected in January 1995, in the Bay of Málaga, North Alboran Sea. L _S, wet mass and DNA, RNA and protein content were closely related allometrically to otolith radius ( R ). Larval daily length increments decreased but DNA, protein and wet mass daily increments increased with larval age. Daily length increments showed a negative and poor relationship with long-term otolith growth. In contrast, DNA, protein and wet mass daily increments were positively correlated. Differences between observed and back-calculated otolith radius-at-age indicated that larvae with slow otolith growth were under represented in older age groups, suggesting the existence of growth-selective mortality. Recent otolith growth, estimated from the mean widths of the last six increments, increased with age and R . Individual RNA: DNA and protein: DNA ratios were correlated significantly, although weakly, with L _S and larval growth.     

长江口鮻矢耳石形态特征及质量与年龄的关系

为研究长江口鮻矢耳石形态特征及其质量与年龄的关系,2017年2—6月从长江口水域采集鮻358尾,解剖并提取耳石324对,并对其形态学参数进行测量分析.结果表明:鮻矢耳石具有基叶和翼叶,中央听沟明显,矢耳石的大小和形状随个体生长变化明显,矢耳石由边缘规则的圆润瓜子状渐变为褊窄的叶片状,边缘波浪状突起逐渐变多.矢耳石平均密度为1.52 mg·mm^-2,平均矩形趋近率为0.68,耳石长占其体长的0.021%~0.047%,耳石宽占其体长的0.009%~0.021%,耳石质量占其体质量的0.045‰~0.731‰.鮻矢耳石长(OL)、宽(OW)、质量(OM)均与体长呈显著相关,耳石宽与耳石质量函数关系式的决定系数(R^2)最高(0.928).耳石质量与体长(BL)最佳拟合方程为幂函数:OM=0.0009BL1.8737(R2=0.967);耳石质量与年龄(A)以及体长与年龄的最佳拟合方程为多项式:OM=2.9262A^2+4.8437A+2.1894(R^2=0.847);BL=-3.2248A^2+102.54A+38.373(R^2=0.858),矢耳石质量与年龄呈显著线性相关,用耳石质量与年龄关系估算的年龄与从耳石上直接读取的年龄无显著差异,矢耳石质量可以作为鉴定鮻年龄的有效辅助手段.     

Rapid larval growth predisposes sex change and sexual size dimorphism in a protogynous hermaphrodite, Parapercis snyderi Jordan & Starks 1905

The temporal relationship between growth history, sex-specific growth divergence and sex change was investigated in the haremic sandperch Parapercis snyderi using otolith microstructure and gonad histology. Parapercis synderi was found to display rapid near-linear growth with a maximum longevity of 303 days. All individuals matured first as female and later changed sex to become male (monandric protogynous hermaphroditism). Individual age-based growth histories obtained from otolith increment widths illustrated that males were larger than females at any given age. Males were found to diverge from the female growth trajectory during two ontogenetic periods; during the larval period and during the period that sex change took place. In addition, male otoliths contained a discontinuity, or 'check mark', associated with the rapid increase in otolith growth during the sex-change period. This microstructural feature was absent from all female otoliths. Accelerated growth in male otoliths lasted up to 25 days, following check-mark formation, after which time otolith growth returned to the pre-check-mark rate. Given the isometric relationship between otolith and somatic growth in P. synderi , and the temporal relationship between the time of check-mark formation and gonad condition, these results strongly suggest that individuals accelerate somatic growth during sex change to become the largest members of the population. Moreover, evidence suggests that the factors that determine the initial growth of larvae influence which individuals will later become males and achieve the highest reproductive success.     

Restricted fish feeding reduces cod otolith opacity

The purpose of this work was to examine the effect of reduced feeding and constant temperature on cod otolith opacity. Three groups of juvenile cod were given restricted food rations at different times for 4 months, resulting in depressed somatic growth. Otolith opacity was measured on pictures of the otolith sections. The otolith carbonate deposited during the experimental period was generally opaque compared to the more translucent otolith material deposited prior to and after the experimental period, when the fish were kept in a pond and in sea‐cages at higher temperatures. Large variations in otolith opacity were found between individual fish both within groups and between groups. In two of the three groups significantly more translucent otolith material was deposited in response to reduced feeding. Our results show that variations in feeding and hence fish growth resulted in variation in otolith opacity, but the effect was minor compared to that of variations in ambient temperature. The combined influence of these effects, which both act on fish metabolism, are most likely controlling the seasonal opacity changes observed in wild fish. Our results help explain the variations seen in fish at constant temperatures.     

Effects of variable ration levels on direct and indirect measures of growth in juvenile red drum (Sciaenops ocellatus)

Relationships between somatic growth (length and weight) and two indirect measures of growth (otolith growth, RNA/DNA ratio) were assessed for red drum (Sciaenops ocellatus) under different feeding rations [0%, 2.5%, 5%, 10%, 15%, and 20% body weight (BW)/day] for 30 days. Representative samples from each ration level were taken in 10-day intervals between Day 0 and Day 30 for evaluation of direct and indirect growth measures. Positive correlations were observed between ration levels, somatic growth, and otolith growth. Statistical differences in weight and length of red drum were observed among ration levels by Days 10 and 20, respectively. Statistical differences for measures of otolith growth among ration levels were evident by Days 20 and 30. In addition, RNA/DNA ratios showed clear separation between fish that were starved and fish that were fed but demonstrated minimal separation among ration levels. Overall, the combination of a measure of somatic growth (weight) and a measure of otolith growth (otolith weight) resulted in the most statistical separation among ration levels. Findings from this study suggest that somatic growth, otolith growth and RNA/DNA ratios are suitable measures of relative growth of red drum; however, due to differences in sensitivity, caution must be exercised when using indirect growth (otolith growth, RNA/DNA ratios) measures to estimate recent growth.     

The record of daily growth in otoliths of Atlantic silversides, Menidia menidia, from field and laboratory

Otoliths were removed from field-collected silversides of age less than 3 months. Otolith diameter was highly correlated with total length of the fish. Daily growth ring counts for this species are known to be a function of age rather than size, so widths for the daily growth rings provide a record of daily increases in length of the fish. Measurement of ring widths showed that weekly specific growth rate was greater than 70% at age 1 week, but declined to about 30% at age 1 month and about 15% at age 2 months. A laboratory experiment in which temperature was changed on a weekly basis demonstrated that environmental variables can affect the width of rings. Nevertheless, the growth rate of field-collected fish, as calculated from otolith ring widths, was more highly correlated with size of fish, as measured by otolith radius, than with the environmental variables of temperature, salinity and plankton abundance. Back-calculation of growth rates from otolith ring widths of five fish collected at the end of the growing season yielded the same age-growth curves as were obtained from 203 fish collected biweekly during the season.     

Otolith accretion rates: Does size really matter?

This study examined the relationship between otolith size and growth in juvenile cod (Gadus morhua L.). Two groups of juvenile cod were reared under different food ration and temperature regimes to obtain fish of similar somatic size but with different sized otoliths. The two groups were subjected to alternating temperature regimes and intermediate ration levels. Large otoliths grew significantly faster than the small ones and variation between individuals was extensive. The ratio of otolith growth during cold and warm temperature exposure did not differ between groups, and the observed growth pattern is therefore not attributable to differential growth within individual temperature periods. The ratio decreased with otolith size, presumably as a result of ontogenetic decrease in otolith protein composition. These results suggest that processes coupled to the metabolic rate of the endolymphatic epithelium are the key driver behind otolith growth.     

Enigmatic ear stones: what we know about the functional role and evolution of fish otoliths

Otoliths in bony fishes play an important role in the senses of balance and hearing. Otolith mass and shape are, among others, likely to be decisive factors influencing otolith motion and thus ear functioning. Yet our knowledge of how exactly these factors influence otolith motion is incomplete. In addition, experimental studies directly investigating the function of otoliths in the inner ear are scarce and yield partly conflicting results. Herein, we discuss questions and hypotheses on how otolith mass and shape, and the relationship between the sensory epithelium and overlying otolith, influence otolith motion. We discuss (i) the state‐of‐the‐art knowledge regarding otolith function, (ii) gaps in knowledge that remain to be filled, and (iii) future approaches that may improve our understanding of the role of otoliths in ear functioning. We further link these functional questions to the evolution of solid teleost otoliths instead of numerous tiny otoconia as found in most other vertebrates. Until now, the selective forces and/or constraints driving the evolution of solid calcareous otoliths and their diversity in shape in teleosts are largely unknown. Based on a data set on the structure of otoliths and otoconia in more than 160 species covering the main vertebrate groups, we present a hypothetical framework for teleost otolith evolution. We suggest that the advent of solid otoliths may have initially been a selectively neutral ‘by‐product’ of other key innovations during teleost evolution. The teleost‐specific genome duplication event may have paved the way for diversification in otolith shape. Otolith shapes may have evolved along with the considerable diversity of, and improvements in, auditory abilities in teleost fishes. However, phenotypic plasticity may also play an important role in the creation of different otolith types, and different portions of the otolith may show different degrees of phenotypic plasticity. Future studies should thus adopt a phylogenetic perspective and apply comparative and methodologically integrative approaches, including fossil otoliths, when investigating otoconia/otolith evolution and their function in the inner ear.     

Somatic and otolith growth in the oyster toadfish (Opsanus tau L.)

Otoliths from oyster toadfish were measured and examined for annuli and daily increments. Distinct annual increments made it feasible to determine growth parameters which demonstrated sexual dimorphism in growth. Microincrements, judged to be daily on the basis of two separate criteria, were enumerated and measured to provide verification of annuli. Utilization of multivariate mathematical models relating age to otolith growth and somatic growth simplified age determination. These methods for examining otoliths should be applicable to other fish species and make it possible to link growth and mortality rates to life history and environmental occurrences.     

Otolith and somatic growth rates in Atlantic salmon parr, Salmo salar L: evidence against coupling

It is often assumed that otolith growth is in some way dependent on somatic growth (i.e. that the two processes are coupled). We examined the relationships between sagitta radius and fork length in 0+ Atlantic salmon parr that would subsequently smolt aged 1 + (UMG fish) or 2+ (LMG fish). Repeated measurements of fork lengths of individually marked parr, taken over a 211-day period from first feeding, were compared to sagitta radii on the same measuring dates (obtained by analysis of daily increments). The results showed that there was a linear relationship between fork length and otolith radius in UMG parr. However, this was not true for LMG parr. These fish enter a state of natural anorexia in their first autumn (despite excess food), but their otoliths continued to grow at the same rate despite the virtual cessation of somatic growth; they had therefore developed disproportionately large otoliths by the end of the study period. The relative growth rates of soma and otoliths first changed in LMG fish in late July/early August; this is the most precise estimate yet obtained of the timing of divergence in the developmental pathways of UMG and LMG parr. The rate of sagitta accretion was consistently lower in LMG parr, possibly indicating a lower metabolic rate in these fish. The results are discussed in relation to previous theories of the relationship between otolith and somatic growth.