Growth dynamics of myotomal muscle fibres in a carangid, Caranx malabaricus (Cuv. et Val.)

The growth pattern of myotomal red, pink and white muscle and its relation to somatic growth in Caranx malabaricus are described. The growth pattern of red muscle was by an increase in fibre number in early size classes (< 22 cm f.l.) and thereafter mainly by increase in fibre diameter and partly by increase in fibre number. The growth of pink muscle was mainly by an increase in fibre diameter, but in smaller fish an increase in fibre number was also evident. White muscle growth was mainly by an increase in fibre diameter and partly by increase in fibre numbers in fish < 22 cm f.l., but only by an increase in fibre diameter from 22 cm f.l. onwards. Caranx malabaricus is a slow-to-moderate growing species and its fibre growth pattern matches with such somatic growth.     

Recruitment and maximal diameter of axial muscle fibres in teleosts and their relationship to somatic growth and ultimate size

Growth of white axial muscle fibres of ten species of freshwater teleosts from five families (Cyprinidae, Centrarchidae, Percidae, Salmonidae, Esocidae) possessing widely different growth rates and ultimate sizes have been studied. The dynamics of muscle increase (i.e. increase in fibre numbers and/or diameter) appears to determine the ability for rapid somatic growth and large ultimate size in teleosts. Thus, the largest and fastest growing species (smallmouth bass, lake whitefish, rainbow trout, muskellunge) show evidence of sustained recruitment of muscle fibres to a large size, in contrast to the smaller and slower growing species (bluntnose minnow, longnose dace). Pumpkinseed, bluegill and yellow perch are all intermediate in fibre growth dynamics, growth and ultimate size between the smaller and larger species. Moreover, the ability of teleosts to grow rapidly and attain a large ultimate size is dependent on the body length at which recruitment of new muscle fibres into the growing axial muscle ceases. The regression equation, y =– 0.29 + 2.26 ( x ), showing the relationship (r = 0.95) between ultimate body length, x , and fork length at cessation of recruitment, y , indicated that for these teleosts, recruitment tends to cease when the fork length reaches about 44% of the ultimate body length. Possible mechanisms to account for this relationship are proposed, and the role of the ultimate fibre diameter in posing limits to the ultimate size of the species is discussed.     

The relationship between mosaic muscle fibres and size in rainbow trout (Salmo gairdneri)

The dynamics of increase of the mosaic muscle in hatchery reared rainbow trout (2·3 to 61·3 cm fork length) are investigated. In trout <5 cm, all fibres are <40 μm in diameter; from 5 to 20 cm the diameter mode remains in the 0–39·9 μm class and there is some extension in range of diameter, thereby suggesting that mosaic muscle increase up to 20 cm is mainly by recruitment of new (small) fibres. When trout exceed 20 cm, mode of fibre diameter shifts to the 40–79·9 um class and fibres of larger diameter (> 100 μrn) appear but the subsequent overall fibre diameter frequency distribution changes little until 50 cm. Increase in muscle during the phase 20–50 cm seems partly attributable to increase in fibre diameter, but remains largely the result of recruitment of small fibres, although the mechanism of the latter process appears less clear than in fish of <20 cm. The recruitment of new fibres stops at 55 cm and further increase in mosaic muscle evidently depends upon the ability of existing fibres to increase in diameter. Possible means by which increasing muscle fibre diameter may limit the ultimate size which trout may achieve are discussed. A phenomenon of apparent reduction in fibre diameter in winter among fish 20–39·9 cm long is noted and its significance is considered.     

Growth dynamics of white and red muscle fibres in fast- and slow-growing strains of rainbow trout

Compared with fish of a slow-growing strain, fast-growing rainbow trout exhibited significantly smaller white fibre diameters, throughout development from hatching to 24 cm body length, although possessing similar total number of fibres. In contrast, in red muscle, no differences were observed in fibre diameter between the two strains, but the fast growing fish showed a significantly higher number of red fibres. The differences in growth rate between the two strains were related to the mean white fibre diameter and were found to be matched by proportional adjustments in recruitment of new fibres to the growing muscle. Thus, the largest and fastestgrowing strain showed evidence of sustained higher recruitment of muscle fibres that endowed this strain with the potential to maintain rapid somatic growth for longer and accomplish greater muscle growth.     

Regional variations in fibre growth dynamics of myotomal and caudal fin muscles in relation to body size of a freshwater teleost, Barbus sarana (Cuv. & Val.)

The growth of red fibres in anterior and middle myotomal regions of B. sarana was mainly by hyperplasia in smaller size classes. In higher size classes, growth by hyperplasia was greater in posterior myotomal region compared to the other two myotomal regions. The growth of pink fibres in anterior myotomal regions was mainly by hypertrophy. The middle and posterior myotomal regions showed fibre growth by hyperplasia. The growth dynamics of white fibres revealed more or less similar pattern in all three myotomal regions against the somatic development. White fibres grew by hyperplasia up to 8 cm F.L. size classes and thereafter by hypertrophy. However, in > 12 cm F.L. size classes, the mean diameter of white fibres did not increase significantly. Similar pattern of growth was found in the white fibres of caudal fin muscle. It is interesting to note that the hyperplasia was mostly completed in the white fibres of the smallest fish studies, whereas, it continued to quite larger fish size in red and pink fibres. Thus, hyperplasia and hypertrophy may be responsible for growth in all fibre types in all myotomal regions in relation to somatic development in this small and medium growing species.     

Growth dynamics of caudal and pectoral fin muscle fibres in a carangid, Caranx malabaricus (Cuv. & Val.), and their possible relation with somatic growth

The growth dynamics of red, pink and white fibres of the caudal and pectoral fin muscles are described in Carans malabaricus (Cuv. & Val.) in relation to their somatic growth. In all three fibre types growth occurred by an increase in fibre number and diameter in small size classes of fish and by an increase in diameter only in larger fish. The growth dynamics of the three fibre types were similar to those of the myotomal muscle fibres and paralleled the somatic growth pattern of this fish.     

Studies on the swimming musculature of the rainbow trout I. Fibre types

A histochemical study has been made on the myotomal musculature of the rainbow trout Salmo gairdneri Richardson. Four main types of fibre can be distinguished on the basis of differences in fibre size, lipid content and succinic dehydrogenase and myofibrillar adenosine triphosphatase activity (ATPase activity). Early histological studies have concluded that in addition to a superficial strip of muscle small diameter 'red' fibres occur throughout the trout myotome. These small diameter fibres occurring in the bulk of the myotome are shown to differ from the superficial fibres with respect to their oxidative metabolism and myofibrillar ATPase. The relevance of this finding to studies on the energetics and swimming efficiency of this species are discussed.     

Influence of bovine growth hormone on the growth dynamics of mosaic muscle in relation to somatic growth of rainbow trout, Salmo gairdneri Richardson

Bovine growth hormone (bGH), administered intramuscularly at 20 μg^-1 every 2 weeks, produced faster somatic growth in rainbow trout fingerlings to approximately 30 cm than in those not receiving the hormone. Both groups were maintained at 12° C, with a photoperiod of 16 : 8 LD, and on ad libitum rations daily. The faster growth was characterized by a controlled increase in input of fibres into the mosaic muscle (which dominates the body tissues by its bulk). It is not known if the enhanced input of fibres is an example of the known ability of the mosaic fibre mass to respond with flexibility and precision to widely differing somatic growth rates, or is a specific consequence of the growth hormone. In other fingerling trout receiving 4-5% daily rations and consequently growing slowly, bGH did not stimulate growth or evidently modify fibre input dynamics. Body dry weight, and condition ( K ), though modified by rations and growth rate, evidently had little effect on muscle fibre growth.     

Postlarval muscle growth in fish: a DNA flow cytometric and morphometric analysis

The mechanism of postlarval fish myotomal growth was investigated in trout (Salmo gairdneri) by means of morphometric and cytofluorometric analysis. The mechanism by which new fibres are added during postlarval growth (hyperplasia) is not fully understood. In histological cross sections these new fibres have a small diameter which give the muscle a "mosaic" appearance. One hypothesis suggested that they could be derived from the proliferative activity of satellite cells. DNA cytofluorometric analysis of nuclei suspensions obtained from trout white myotomal muscle during different developmental stages (eleutherembyronic; alevin; yearling and adult) showed a consistently low S-cytometric phase during all stage in which myofibres of small diameters were present. The percentage of such small fibres, determined by morphometric analysis, suggested that satellite cells are the proliferative population. In fact, their percentages, as determined by morphometric analysis in histological section, bear a linear relationship with the S-cytometric phase percent nuclei (R = 0.927). Only in adults (67 cm in size) there was a significant decrease in the S-cytometric phase. At this stage, in histological sections, the myotomal muscle no longer had a "mosaic" appearance because of the disappearance of the small fibres. It may, therefore, be supposed that in the cm 67 adult specimens, the proliferative population is entering the G0 phase. It is known, in fact, that muscle growth proceeds only by fibre hypertrophy in trout longer than 70 cm in length (Stickland, 1983).     

Sexual differences in the increase of white muscle fibres in Argentine hake, Merluccius hubbsi, from the San Matias Gulf (Argentina)

Growth dynamics of white fibres from axial muscle has been investigated in Argentine hake, Merluccius hubbsi , discriminating between sexes for the first time. The frequency distributions of fibre diameters are remarkably different in both sexes at sizes between 42 and 63·9 cm total length (T.l.). Males have a much lower proportion of newly recruited fibres (0–10 μm) than females; at 52–53·9 cm T.l., females have 4% of fibres in that category and males 0·5%. It appears that, from this size interval onwards, the increase in muscle mass is due only to the increase in diameter of individual fibres, which may exceed 300 μm. The lower recruitment rate of new fibres in males, and its relationship to lower growth rates and smaller final sizes, are discussed, and possible effects of reproductive activity are considered.     

Hyperplasia and hypertrophy in the growth of skeletal muscle in juvenile Atlantic salmon, Salmo salar L.

Both red and white muscle fibre numbers in juvenile Atlantic salmon increased gradually with fish length throughout the freshwater growth period. Mean fibre area increased as fish grew to 6.5 cm f.l. , but thereafter was unrelated to fish length. Hyperplasia was most obvious when fish were growing fastest, and was the dominant growth process in fish over 6.5 cm f.l. Hypertrophy was most important when growth was slow, as in autumn and winter.
Mean white fibre area was significantly smaller in deep muscle than at medial and superficial sites. Total cross-sectional area of red, white and total trunk muscle increased with fish length. The ratio of red: white cross-sectional area increased with fish length to a plateau at about 10% after 6.5 cm f.l.     

A histochemical study of the lateral muscles of five teleost species

A qualitative histochemical study has been made of the myotomal muscles of five teleost fish (glass fish, Chanda ranga; carp, Carassius carassius; coalfish, Gadus virens; black mollie, Molliensia sp. and grey mullet, Mugil cephalus ) . Three or four main fibre types were distinguished in these species on the basis of the distribution and relative activities of glycogen, lipid, aglycerophosphate dehydrogenase, phosphorylase, and succinic dehydrogenase. The so-called red and white fibre types were found to have similar histochemical properties to previously investigated species. All the species studied, with the exception of the glass fish, Chanda ranga , were found to have one or two types of pink fibre situated between the red and white fibre regions. In the carp, coalfish and mullet, the pink fibres were found to be composed of small and large diameter fibres which were similar to red and white fibres respectively, except for their staining for succinic dehydrogenase. Considerable differences were found in the relative amounts of pink muscles between species. Minor fibre components were found in several species. These consisted of very small diameter fibres which did not stain well with any of the histochemical procedures used. It is suggested that these fibres represent areas of continuing muscle growth. The results obtained are discussed in relation to the division of labour between myotomal muscles during swimming.     

Proliferative activity and motoneurone recruitment persist at the spinal cord central canal during larval and some postlarval stages in the rainbow trout (Oncorhynchus mykiss)

We previously found a linear relationship between the cross sectional myotomal area and the motoneurone number in the growing trout during postlarval stages. These neurones increased in number until a fish length of 150 mm, which prompted us to examine how motor neurones are recruited afterwards to meet the growth of their target myotomal muscle. Young adult (260 mm in length), fingerlings (F, 120-170 mm), fry (Fr, 70 mm) and eleutherembryos (Es, 20-30 mm) of rainbow trout (Oncorhyncus mykiss) were employed in this study. PCNA immunohistochemistry was used for monitoring the proliferative activity in the epithelium of the spinal cord central canal. This activity was quantified as the number of PCNA labelled cells for each spinal cord section. In Es and Fry, a mean value of 3-5 labelled cells for each section was found with a sharp decrease in young F (120 mm long). After this fish length, it was not possible to quantitatively evaluate the proliferative activity at the central canal. However, labelled cells were seldom found in the spinal cord sections until a fish length of 260 mm. From these data it is possible to conclude that motoneurone recruitment in the trout spinal cord is down-regulated at the F stage. Afterwards, we found that motoneurones increase in size to meet the growth of their target myotomal muscle.     

Histochemical characterization of myotomal muscle in the bluntnose minnow, Pimephales notatus Rafinesque

A histochemical study of the axial muscles in the bluntnose minnow, Pimephales notatus Rafinesque, revealed three main types of fibres-red, white and pink. These were distin- guished on the basis of glycogen or lipid contents, or of NADH diaphorase and actomyosin ATPase activities. White muscle formed the largest fraction of the total, whereas red ranged from slightly > 1% of the total cross-sectional area in the immediately post-pectoral fin region to slightly <4% in the post-anal region. Pink muscle formed only a few layers of fibres between red and white. Results are discussed in relation to possible division of labour among the different types of fibres. However, it is emphasized that fibre types may be more functionally versatile in fish muscles than has frequently been supposed.     

Dual motor innervation in the axial musculature of fishes

A systematic survey of motor innervation in the myotomal muscle fibres in several groups of fishes was conducted to observe the extent of dual innervation, a phenomenon found within those myotomal muscle fibres with terminal innervation. It is concluded that the dually innervated myotomal muscle fibre is a primitive vertebrate feature. Furthermore, it is suggested that this particular form of polyneuronal innervation is retained in the course of neural evolution within the homogeneous white myotomal muscle fibre region in these fish groups in order to insure synchrony of firing during sudden, rapid locomotion. The findings are examined in light of current ideas regarding the direct adrenergic innervation in addition to cholinergic innervation of striated muscle fibres.     

Scaling effects on the neuromuscular system,twitch kinetics and morphometrics of the cod,Gadus morhua

Size related changes in muscle twitch kinetics, morphometrics and innervation have been examined in cod, Gadus morhua. Fish size ranged from 9 cm to 45 cm in total length (L).

Twitch contraction time (time to 90% relaxation), scaled in proportion to L^0.29. Scaling of morphometric parameters was essentially geometric. Mean cross‐sectional area and weight of the myotomal muscle scaled in proportion to L^2.05 and L^3.08 respectively. These results are discussed in the light of alterations in length specific swimming performance and kinematics as fish grow.

During growth, the number of endplates terminating on each fast fibre increased, from around 10 on fibres 2 mm in length (～10 cm fish) to 20 on 10 mm fibres (—40 cm fish). However, mean spacing between endplates increased from around 0.25 mm to 0.50 mm. The functional significance of polyneuronal innervation in teleost fast muscle fibres is discussed.     

Histochemical characterization of myotomal muscle in the grass pickerel, Esox americanus vermiculatus (LeSeuer), and the muskellunge, E. masquinongy (Mitchell)

A histochemical study of the myotomal muscles in the grass pickerel, Esox americanus vermiculatus , and the muskellunge, E. masquinongy , was performed using actomyosin ATPase and NADH diaphorase activities. Three fibre types, i.e., red, white and pink were distinguished on the basis of their enzyme activities. White muscle fibres comprised the bulk of the myotomal musculature. The relative proportion of red muscle fibres was greater in the caudal region than in more anterior regions of the body. Pink fibres formed only a few layers between red and white. These findings are discussed in relation to the possible functional significance of the muscle fibre types in swimming and feeding behaviour in these species.     

Influence of growth rate on white muscle dynamics in rainbow trout and brook trout

In trout, fast growth stimulated white muscle fibre hypertrophy ( P 0·001) and hyperplasia ( P <0·01) in outer fibres but not in deep fibres. Glycogen was most prevalent in outer fibres ( P <0·01) and in brook trout ( P <0·01) that on average had three to four times larger fibres than rainbow trout.     

Protein, lipid and caloric contents of bluntnose minnow, Pimephales notatus Rafinesque, during growth at different temperatures

Changes in the relative proportions of protein, lipid, water and caloric contents of bluntnose minnow growing at various temperatures (15, 25, 30° C) were investigated by application of the allometry equation, y=ax^b . Fish grew significantly faster at 25° C (closest to optimum), more slowly at 30° C and most slowly at 15° C. Protein, as a percentage of body wet weight, tended to increase with fish size at all temperatures ( b > 1.000), whereas in juveniles (<0.7 g) it decreased ( b < 1.000). However, with the exception of the 15° C group, protein as a percentage of body dry weight, decreased in all groups ( b < 1.000). Temperature appeared to modify the body composition of bluntnose minnows, e.g. decreasing temperature led to significantly enhanced protein content during growth. Lipid (%) and caloric content (cal g⁻¹) increased with increasing fish weight ( b > 1). The slower growing fish (15°, 30° C) deposited significantly more lipid (and had higher caloric contents) than those growing most rapidly (at 25° C). Water content (%) decreased with increasing body weight in all groups. Despite intergroup growth rate differences, all groups showed evidence of a tendency to follow similar trends in b values for body constituents and caloric content (except for protein v. body dry weight for the 15° C group). This suggests a general conservativeness of body composition in bluntnose minnow. The correlations between body constituents, caloric content and body weight were high ( r ²>0.9) so that estimates of body composition can be obtained from body weight for all temperature groups.     

A morphometric analysis of the spinal motor pool in relation to its target muscle during growth in the trout, Salmo gairdneri Richardson

The relation between number and size of spinal motoneurons and the dimension of myotomal muscle has been investigated in trout at different stages of embryonic, larval and postlarval development (body length 1–15 cm). Three spinal segments have been analysed (cervical, trunk and caudal) and the following parameters were determined by means of a Micromeasurements Image Analyzer: (a) mean cross-sectional myotomal area; (b) mean soma size of principal (or dorsomedian, DM) and secondary (or ventrolateral, VL) motoneurons; (c) DM and VL motoneuron density per segment. Myotomal muscle and motor pool growth was evaluated by percent increments of a, b and c parameters at each stage. Their relationships were denned by equations of computed regression lines.
The analysis provided evidence that: (1) a continuous exponential growth of mean myotomal area takes place in the three segments, with the same trend and with the lowest values in the caudal segment; (2) DM and VL motoneuron size and density per segment also increase during development, with the least value in the caudal segment, VL parameters being of lesser value than DM; (3) motoneuron pool and its target myotomal muscle parameters bear a linear relationship as defined by equations of computer regression lines; (4) motoneuron number percent increment at early eleutherembryonic stage precedes myotomal area increment which takes place during late eleutherembryonic stage.
It is apparent that spinal motor pool and target myotomal muscle grow at the same rate in the trout during the considered stages. The discussion links this fact with the hypothesis of a neuronal influence on muscle fibre type differentiation.