Moult cycle and its chronology in Mysis mixta and Neomysis integer (Crustacea, Mysidacea): implications for growth assessment

The moult-staging technique was used to determine the main moult stages (MS) and their duration in laboratory-reared juveniles and subadults of Mysis mixta and Neomysis integer. The relative duration of each stage was similar for both species with premoult occupying the major part of the moult cycle (51% and 44% for M. mixta and N. integer, respectively), followed by postmoult (26% and 34%) and intermoult (23% and 22%). Effects of temperature and feeding regimes on the chronology of the moult cycle were investigated. When the duration of the moult cycle (MCD) was extended by manipulating the feeding regime, animals prolonged their late postmoult and early premoult stages. At 12 and 5 °C, no specific moult stage varied in relative duration as long as food supply was high. Field application of moult staging for growth assessment was tested using wild-caught M. mixta. The MCD estimated via moulting experiments was compared to that obtained by analyzing moult stage distribution combined with the experimentally obtained data on stage duration. Close correspondence between two methods of MCD assessment was observed with moult-staging technique being particularly useful in situations when conducting of experiments immediately after collection is not feasible.     

Tissue accumulation and the moult cycle in juveniles of the Australian freshwater crayfish Cherax destructor

1. This paper investigates moult stage and size-specific changes in whole body composition during growth in juvenile crayfish in order to better describe the nature of growth and energy use. 2. Composition is described in terms of moult stage and size-specific wet, dry and ash-free dry weight, water, carbon, protein nitrogen, non-protein nitrogen, ash and energy. Dry weight and ash-free dry weight (AFDW) peaked in the middle and in the later stages of premoult. Both peaks were about 2–2.3 times postmoult weight. Losses in tissue weight during ecdysis were substantial in the smallest crayfish but declined with size. Water was taken up between late premoult and early postmoult. Tissue accumulation occurred primarily between B and D₁.₃, with further weight gain largely the result of fluid uptake. Ash increased immediately postmoult with a major peak occurring during intermoult. Mean organic carbon varied between 33 and 35.5% of the body and 49% of the exuvia. Chitin varied between 9 and 17% of body AFDW and made up about 50.5% of the exuvia. Protein content varied between 47 and 62% of body AFDW and about 25% of the exuvia. Carbon, chitin and protein were not affected by moult stage but protein declined with ocular carapace length (OCL) in larger crayfish. Mean caloric content varied between 19 and 22 J mg-¹ AFDW depending on size and moult stage. Caloric content increased with OCL during premoult and early intermoult then declined with size until part of the way through premoult. 3. Relationships between protein, chitin and remaining carbon (organic carbon minus chitin) were examined. It is suggested that protein and some carbon are catabolized during the moulting process, possibly to fuel metabolism. Models are presented showing changes in proximate composition over the moult cycle for two sizes of crayfish, and tissue and energy accumulation and loss over a series of moult cycles and sizes from 3.1 mm to 17 mm OCL.     

Population biology and assessment of the white-spotted spinefoot, Siganus canaliculatus (Park, 1797), in the southern Arabian Gulf

The age, growth, mortality, reproduction and resource status of Siganus canaliculatus in the southern Arabian Gulf were investigated using a combination of size frequency, biological and size‐at‐age data. Defined structural increments consisting of alternating translucent and opaque bands in transverse sections of sagittal otoliths were validated as annuli. The maximum absolute age estimate was 7.8 years. Parameter values of the von Bertalanffy growth function fit to size‐at‐age data (males and females combined) were: k = 1.0, L_∞ = 24.8 cm (L_F), t_o = −0.1 years. Fish in spawning condition were only observed between April and July although patterns in gonadosomatic indices suggested a second but less well defined spawning event in November. The mean sizes and ages at first sexual maturity were 21.5 cm L_F (1.9 years) for males and 25.7 cm L_F (2.1 years) for females. Fish were fully recruited to the fishery at a size (L₁₀₀ = 19.7 cm L_F) that was smaller than the sizes at which sexual maturity was attained. The annual instantaneous rate of fishing mortality (F = 0.85 year⁻¹) (0.26–1.44 year⁻¹ 95% CI) was considerably greater than the target (F_opt = 0.33 year⁻¹) and limit (F_limit = 0.44 year⁻¹) biological reference points, indicating that the stock is overexploited.     

Respiration and growth of Hyas araneus L. Larvae (Decapoda:Majidae) from hatching to metamorphosis

Rates of respiration and growth were measured for larvae of the spider crab Hyas araneus L., reared in the laboratory from hatching to metamorphosis. The moulting cycle was simultaneously monitored. In both zoeal instars individual respiration rate (R) increased as a linear function of time (t) of development, whereas growth, measured as dry weight (W), carbon (C), nitrogen (N), hydrogen (H), and energy content (E, calculated from C) followed a power function of t. Weight-specific respiration rate (QO₂) was in all instars maximum in early postmoult, and minimum in intermoult and early premoult. Zoea II and megalopa instars showed another conspicuous QO₂ increase during late premoult. Respiration (both R and QO₂)and growth of the megalopa could be described by non-linear (quadratic) functions of t. R and QO₂ during this larval stage were not correlated with W, but were controlled by events of the moulting cycle: R followed a similar pattern to QO₂ (minimum values in intermoult), whereas biomass of the megalopa changed conversely, with a maximum in intermoult and early premoult. The respiratory coefficient (i.e. the ratio of metabolic energy loss: energy gain by body growth) was far lower (<0.8) in the zoeal instars than in the megalopa (>5), suggesting a strongly reduced capability of energy conversion in the final larval stage of H. araneus.     

Some biological parameters of the Mediterranean sand smelt Atherina (Atherina) hepsetus, Linnaeus, 1758 (Pisces: Atherinidae) from the middle eastern Adriatic (Croatian coast)

Data on the length–weight relationship, age, growth, sex ratio and mortality were analysed for the Mediterranean sand smelt, Atherina (Atherina) hepsetus L. (total = 2805; males = 1258; females = 1547) collected in the eastern middle Adriatic island area during the reproductive period (February to April) in 2002. The total length of sampled specimens ranged from 3.8 to 14.5 cm and the weight from 0.28 to 22.39 g. The overall sex ratio was 1.23 : 1 in favour of females, significantly different from the expected 1 : 1 ratio (χ² = 29.76; P < 0.05). All individuals >13.4 cm were females. The oldest collected male and female specimens were 5 years old. The von Bertalanffy growth formula was estimated for females (L_∞ = 15.79 (1−e^{−0.43(t+0.049)}) and males (L_∞ = 15.25 (1−e^{−0.43(t+0.018)}). The power values (b) of the length–weight relationship were very similar for both sexes (b = 3.14) and indicated a slightly allometric growth. The instantaneous rates of mortality for all collected fish were Z = 1.44 year⁻¹; M = 0.94 year⁻¹ and F = 0.50 year⁻¹. The exploitation ratio was E = F/Z = 0.35. The value for M is highly uncertain, however, as well as those values for F and E.     

Moult cycle and morphogenesis inHyas araneus larvae (decapoda,majidae), reared in the laboratory

Moult cycle and morphogenesis in larval instars (zoea I, zoea II, megalopa) of the spider crabHyas araneus (L.) were studied in the laboratory. Changes in the epidermis and cuticle were documented photographically at daily intervals to characterize the stages of the moult cycle. Stage A (early postmoult) is a very short period during which the larva takes up water. During late postmoult (B) and intermoult (C) the endocuticle is secreted, and there is conspicuous epidermal tissue condensation and growth. The onset of early premoult (D₀) is characterized by epidermal apolysis, occurring first at the bases of the setae in the telson of zoeal instars or in the rostrum of the megalopa, respectively. Intermediate premoult (D₁) is the main period of morphogenesis, in particular of setogenesis: in the setae of the zoeal telson and carapace there is invagination or (in the zoea II) degeneration of epidermal tissues. Formation of new setae in the interior of epidermal tubules was observed in zoeal maxillipeds and in the antennae of the zoea II and megalopa instars. During late premoult (Stages D_2–4) part of the new cuticle is secreted, and the results of morphogenesis become clearly visible. For technical reasons (rigid exoskeleton) only a preliminary account of the moult cycle in the megalopa can be given. A time schedule is suggested for the stages of the moult cycle. It is estimated that postmoult (A–B) takes ca 9 to 15 % of total instar duration, intermoult (C) ca 22 to 37 %, and premoult (D) ca 48 to 69 %. There is an increasing trend of relative portions of time (% of total instar duration) from instar to instar in Stages A–C (mainly in the latter) and a decreasing trend in Stage D (mainly in D₀ and D_2–4).     

Age, growth and reproductive characteristics of chub, Leuciscus cephalus (L., 1758) in the İkizcetepeler dam lake (Balikesir), Turkey

In this study, the population structure, growth and reproduction characteristics of 414 chub (Leuciscus cephalus L., 1758) from the ?kizcetepeler dam lake were investigated monthly between January and December 2000. Age groups ranged between I and VI for this species in the reservoir, with the second and third year‐classes dominating. Sex ratio was 1 : 1.4 (M : F), corresponding to 58.4% males and 41.6% females. Females attained greater size and age than males. The largest female captured was 24.8 cm FL, the largest male was 24.1 cm FL, both age VI. The von Bertalanffy growth equations and length–weight relationships were found as: L_t = 28.89[1 ?e^{?0.224(t+1.55)}] for females, L_t = 26.71[1 ? e^{?0.259(t+1.55)}] for males; W_t = 347.386[1?e^{?0.224h (t+1.55)}]^2.86 for females, W_t =286.48[1?e^{?0.259 (t+1.55)}]^2.92 for males; W = 0.0227 × L^2.87 for females and W = 0.0194 × L^2.92 for males. Significant statistical differences in condition factors between age classes and sexes were not found (P > 0.05, t‐test). Spawning period of this species in the lake was between April and May.     

Osmoregulatory capacity,health status and growth as functions of moult stages from various weight classes in marron (Cherax cainii) and yabbies (Cherax destructor)

The study investigated the relationship between five moult stages and the osmoregulatory capacities (OC), moisture content of hepatopancreas (HM%), hepatosomatic indices (HI_wet, HI_dry) and growth of 6 weight classes of 235 marron (Cherax cainii) and 235 yabbies (Cherax destructor) under laboratory conditions. In each moult stage, OC increased linearly with the increase in wet body weight. Intermoult stage C showed the highest OC significantly lower than premoult stages D₀, D₁, D₂ and postmoult stage AB in every weight class in both crayfish. HM% decreased from AB to D₂, while both HI_wet and HI_dry reduced from C to D₂. Percentage dry matter of whole body carcass was highest in stage C and lowest during the AB stage with a significant difference between the two species. Larger than 15-g marron and yabbies differ with each other in OC, HI_wet, and HI_dry, moult intervals and growth rates.     

Calculating growth parameters of Genidens barbus (Siluriformes, Ariidae) using length composition and age data

The marine catfish Genidens barbus was once one of the most important estuarine fishery resources in the Patos Lagoon (southern Brazil), which had the largest population known for the species. However, clear signs of overfishing have been observed since the 1980s. Growth parameters for this population were estimated by combining length frequency data for juveniles and selected published data on annuli reading from adult otoliths. This methodology yielded a more realistic set of parameters using both the specialized and generalized von Bertalanffy growth model. The species growth can be described as: L_t = 118.6[1 − e^{−0.043(t+1.505)}] or L_t = 104.6[1 −e^{−0.137D(t+4.013)}]^1/D with the surface factor D = 0.542. The growth performance index ϕ′ was estimated as 2.78 for the special (classical) model. Stock rebuilding is expected to be slow, as the species is slow‐growing, has a low natural mortality rate, and mouthbreeds a small amount of large eggs.     

Modeling population dynamics of yeast-like symbionts (Ascomycota: Pyrenomycetes: Clavicipitaceae) of the planthopper <Emphasis Type="Italic">Delphacodes kuscheli</Emphasis> (Hemiptera: Delphacidae)

Delphacodes kuscheli establish mutualistic relationship with yeast-like symbionts (YLS) that live in the fat body and are necessary for host survival and reproduction. We estimated for a host of age t, its body weight, W_(t), and the number of YLS per host, YLS_(t). The host body weight was calculated as: W_(t)?=?Lm/[1+ e ^(d–kt)], (Lm?=?the maximum observed weight, and d and k are constants), and the fat body was considered a fixed proportion of W_(t). We calculated the number of YLS per unit host body mass: α_(t)?=?YLS_(t)/W_(t). We also calculated the number of YLS per host, cYLS_(t), and analyzed the pattern of variation in both sexes adapting the expression of the logistic model: cYLS_(t)?=?KN_oe^rt/K+(e^rt -1)N_o, (N_o?=?initial number of YLS, r?=?intrinsic per capita rate of natural increase, and K?=?variable carrying capacity). In females the carrying capacity varied according to a constant proportion of the host’s weight: K_(t)?=?αW_(t). In males α_(t) was considered a decreasing function of the host age: K_(t)?=?α_(t)W_(t). The coefficients No, α, and r were subjected to parameterization. We found that the patterns of W_(t) and YLS_(t) of D. kuscheli were similar to other planthoppers. In females YLS increased up to the adult stage and then remained almost constant, varying similarly to individual weight. In males YLS increased up to the 5th instar nymph as the individual weight did, but the number of YLS decreased in the adult stage and the correlation was not so good. The calculated number of YLS per host matches reasonably well with the number estimated experimentally both in females and males. This is the first study that quantified and modeled the dynamics of YLS endosymbionts in a Neotropical planthopper pest. The models will be used in future studies for better understand the experimental reduction of YLS in young nymphal stages.     

Haemolymph constituents and osmolality as functions of moult stage,body weight,and feeding status in marron,Cherax cainii () and yabbies,Cherax destructor (Clark, 1936)

The study investigates the change in osmolality and haemolymph constituents in marron Cherax cainii and yabbies Cherax destructor associated with moult stages, body weights and their feeding status. A total of 582 haemolymph samples from 5 moult stages (postmoult-AB, intermoult-C, and premoult stages – D₀, D₁, D₂), two body weight classes (2–15 g and 61–75 g) and nutritional status were used for analysis of osmolality, protein, glucose, and ionic concentrations of potassium and chloride following the standard biochemical procedures. The haemolymph protein, glucose, potassium and chloride levels were highest at intermoult and early premoult stages, and lowest at postmoult in both crayfish species. Except protein, no significant differences were seen in analyzed parameters between various weight classes and two species. Haemolymph osmolality, protein and glucose were significantly higher in fed crayfish, whereas no variations in haemolymph potassium and chloride concentrations were observed between the fed and unfed crayfish. Maximum osmolality was recorded at 7–8 h after feeding in both crayfish species. The results showed that the biochemical changes in the haemolymph of marron and yabbies are related to moult stages, body weight and feeding and thus can be used as tools for determining suitable diets.     

A sexual conflict in collared flycatchers,Ficedula albicollis: early male moult reduces female fitness

A sexual conflict over levels of parental care occurs in most animals with biparental care, and studies of sexual differences in levels of parental care have usually focused on its intra-annual fitness consequences. We investigated inter-annual fitness consequences of a sexual difference in timing of feather replacement (moult) in collared flycatchers (Ficedula albicollis). In this study, males overlapped reproduction and moult more often than females, they also initiated their moult at an earlier stage of breeding than females. Females mated to males with a moult-breeding overlap had significantly lowered survival chances than females mated with males initiating moult after breeding. Furthermore, females mated with moulting males risked a lowered future fecundity in terms of a delayed start to breeding in the following season. However, early moulting males achieved a similar reproductive success as males initiating moult after breeding. Likewise, male survival probability to the following breeding season did not differ between early and late moulting individuals, nor was there any evidence that males gained or lost in future mating advantages by moulting early. These results show not only that a sexual conflict over timing of moult may operate, but also that it can impose severe fitness consequences, in terms of reduced future fecundity and survival probability, upon the ''losing'' sex.     

Age and growth of Gerres oyena (Forsskål, 1775) on Okinawa Island, Japan

Age and growth were studied for Gerres oyena (Forsskål, 1775) on Okinawa Island, southern Japan from November 2002 to November 2005. A total of 408 samples was collected ranging from 5.85 to 19.65 cm standard length (L_SL). Male fish age was estimated at up to 6⁺ years, whereas females reached 8⁺ years as estimated by sectioned otoliths. The length–weight relationships and the von Bertalanffy growth curve were described for all individuals as: and L_t = 20.54{1 − e^{−0.1807(t+2.8462)}}, respectively. Opaque rings were formed from April to August during the spring–summer seasons.     

Life history of the river goby Glossogobius callidus (Teleostei: Gobiidae)

The river goby Glossogobius callidus is native to freshwater and estuarine habitats in South Africa. Individuals [21.1–144.4 mm total length (L_T)] were sampled from impoundments in the Sundays River Valley, Eastern Cape, from February 2014 to March 2015. The largest female was 137.2 mm L_T, and the largest male was 144.4 mm L_T. Length-at-50% maturity was 75.2 ± 2.1 mm L_T for males and 76.2 ± 2.0 mm L_T for females. Absolute fecundity was 1028.2 ± 131.7 oocytes per fish, and relative fecundity was 50.1 ± 18.1 oocytes per gram. The spawning season extended from October to December. Fish were aged using sectioned sagittal otoliths. The growth zone periodicity was validated using edge analysis. Longevity was more than 7 years for females and more than 6 years for males. Length-at-age was similar for the two sexes and was best described using the von Bertalanffy growth model as L_t = 74.7(1 − e^{–1.0(t + 0.1)}) mm L_T for the entire population. Using the population age structure, the mortality rate was estimated at 1.3 per year.     

Age,growth and reproduction of the sand smelt Atherina boyeri Risso, 1810 in Mellah Lagoon (Eastern Algeria)

This aim of this paper was the study of the reproductive biology and growth of the sand smelt, Atherina boyeri, in Mellah Lagoon (Algeria). These data are important for the sustainable exploitation of the stocks of this species. Examined was a total of 1402 Atherina boyeri specimens captured monthly from March 2010 to March 2011, in a population with a 3‐year life cycle. Length–weight relationship was estimated as W = 0.0047 L^3.077 (r² = 0.935) for males and W = 0.0047 L^3.176 (r² = 0.935) for females. Using scales, the von Bertalanffy growth function fitted to back‐calculated size‐at‐age data was Lt = 9.49 [1 ? e^{?0.316 (t + 0.928)}] for males, and Lt = 11.67 [1 ? e^{?0.179 (t + 1.514)}] for females; using otoliths this was Lt = 9.68 [1 ? e^{?0.3 (t + 1.02)}] for males, and Lt = 11.93 [1 ? e^{?0.171 (t + 1.55)}] for females. The growth performance index (Φ) indicated that males (Φ_scales = 3.34, Φ_otoliths = 3.33) grew at the same rate as females (Φ_scales = 3.19, Φ_otoliths = 3.24), with a sex ratio of 1 : 1.6 in favor of females. The reproductive season extended from February to June. Individual length at first sexual maturity was 4.20 cm for 1‐year‐old males and 4.35 cm for 1‐year‐old females.     

Effect of bilateral eyestalk ablation on ovarian development and moulting in early and late intermoult stages of female spiny lobster Panulirus homarus (Linnaeus, 1758)

Abstract

The present study describes the effect of bilateral eyestalk ablation (BESA) on reproduction and moulting of spiny lobster Panulirus homarus females in their early and late intermoult stages. The lobsters obtained from the wild were conditioned for the experiment. The experiments were conducted at the Calicut research centre of Central Marine Fisheries Research Institute, India. Eyestalk ablation of females was done by ligation. The responses of the bilaterally eyestalk-ablated lobsters were statistically analysed. BESA conducted on the early and late intermoult stages resulted in the simultaneous acceleration of the somatic growth and reproductive processes with higher emphasis for oogenesis in lobsters ablated in the early intermoult phase and comparatively lower activity in those ablated in the late intermoult phase. Ablation in late intermoult phase resulted in faster entry into the premoult stage compared to the control.     

Evaluating bony structures for ageing and growth parameters of Capoeta banarescui inhabiting the lower Melet River (Ordu,Turkey)

The aim of this study was to investigate the reliability of bony structures for age determination. A total of 247 Capoeta banarescui individuals were analyzed. Fish were caught on the lower Melet River between July 2010 and October 2012. Sex ratio was 2.5 : 1. The specimens ranged from 8.1 to 21 cm in total length and 5.07 to 108.93 g in weight. Age determined from vertebrae, scales, asteriscus and lapillus otoliths were variable. The highest percent agreement (PA) (69.2%) and the lowest average percent error (APE) (4.24%) and coefficient of variation (CV) (8.71%) were obtained for vertebrae. Hence, it was determined that the vertebra is the most reliable ageing structure for C. banarescui in the lower Melet River. Females were age I–V years and males I–V years. The von Bertalanffy growth equation was established for females, males, and all individuals, respectively: L_t = 23.9 [1?e^{?0.22 (t+1.89)}], L_t = 21.8 [1?e^{?0.26 (t+1.58)}], L_t = 26.7 [1?e^{?0.17 (t+2.21)}].     

Life history of the red-banded seabream Pagrus auriga (Sparidae) from the coasts of the Canarian archipelago

Red‐banded seabream Pagrus auriga (N = 615) were caught off the Canary Islands from January 2003 to December 2004. Total length ranged from 120 to 780 mm. The species was characterized by protogynous hermaphroditism. The male :female ratio was in favour of females (1 : 8.2). The reproductive season extended from September to February, with a peak in spawning activity in October–November. Fifty percentage maturity was reached at 387 mm total length by females and 533 mm by males. The length–weight relationship for all individuals was described by the parameters: a = 0.0086 and b = 3.014, when length is given in mm and weight in grams. Otolith age readings indicated that the population consists of 19 age groups, including a very high proportion of individuals between 0 and 7 years old. Growth analysis reveals that the species is slow‐growing and relatively long lived (18 years). The von Bertalanffy growth parameters for the entire population were: L_∞ = 803 mm, k = 0.081 year⁻¹ and t₀ = −2.17 year. Growth differed between males and females. The instantaneous rate of natural mortality for all fish was: M = 0.164 year⁻¹.     

Age and growth of the blackchin guitarfish Glaucostegus cemiculus (Geoffroy Saint-Hilaire, 1817) from Iskenderun Bay (Northeastern Mediterranean)

Between September 2010 and June 2012, a total 291 (166 females and 125 males) blackchin guitarfish Glaucostegus cemiculus were captured by a commercial bottom trawler (F/V Coşkun Reis) in Iskenderun Bay, Turkey (northeastern Mediterranean Sea). The total length (L) and total weight (W) of the female and male guitarfish ranged between 32.0–165.0 cm and 88 g–16.68 kg, and 34.3–128.3 cm and 112 g–6.00 kg, respectively. Vertebral age estimates ranged from 0 to 8 years for females and 0 to 5 years for males. The growth models of von Bertanlanffy and Gompertz were fitted to the length at age data using the nonlinear regression method. Model selection was based on the values of the residual standard error and the Akaike's information criterion corrected for small sample size (AIC_C) associated with each fit. The von Bertalanffy growth model provided the best fitting growth curves for each sex with parameters reaching L_∞ = 187.17 cm, K = 0.195 year^-1, t₀ = −1.38 year for females, and L_∞ = 144.85 cm, K = 0.321 year^-1, t₀ = −1.13 year^-1 for males. The W–L relationship parameters did not differ significantly between sexes, the estimated values of a and b were 0.0018 and 3.11, respectively. By using these values of a and b, and also respective estimates of L_∞, the values of W_∞ were obtained as 20.53 kg for females and 9.25 kg for males. The overall percentage ratios of females and males in the samples were 57% and 43% respectively.     

Thermodynamics of hexokinase-catalyzed reactions.

The enthalpies of the hexokinase-catalyzed phosphorylation or glucose, mannose, and fructose by ATP to the respective hexose 6-phosphates have been measured calorimetrically in TRIS/TRIS HCl buffer at 25.0, 28.5, and 32.0°C. The effects on the measured enthalpy of the glucose/hexokinase reaction due to variation of pH (over the range 6.7 to 9.0) and ionic strength (over the range 0.02 to 0.25) have been examined. Correction for enthalpy of buffer protonation leads to δH^o and δC_p^o values for the processes: eq-D-hexose + ATP⁴⁻ = eq-D-hexose 6-phosphate²⁻ + ADP³⁻+ H⁺. Results are δH^o = −23.8 ± 0.7 kJ · mol⁻¹ and δC_p^o = −156 ± 280 J·mol⁻¹·K⁻¹ for glucose. δH^o = −21.9 ± 0.7 kJ·mol⁻¹ and δC_p^o = 10 ± 140 J·mol⁻¹·K⁻¹ for mannose, and δH^o = −15.0 ± 0.9 kJ·mol⁻¹ and δC_p^o = −41 ± 160 J·mol⁻¹·K⁻¹ for fructose. Combination of these measured enthalpies with Gibbs energy data for hydrolysis of ATP⁴⁻ and that for the hexose 6-phosphates lead to δS^o values for the above hexokinase-catalyzed reactions.