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1.
Like all poikilotherms, the growth and reproduction of blue crab, Callinectes sapidus depends on temperature and season. Warmer water temperatures in the Chesapeake Bay allow for ovarian development and spawning, while colder water temperatures slow their metabolism and reproduction. The current study aimed to identify optimal environmental conditions for inducing reproduction in animals held in long‐term captivity for year round production in aquaculture through environmental manipulations. Temperature and photoperiod were the main environmental factors tested for 25 weeks: 11°C and 21°C, with the following photoperiods: 0L:24D, 8L:16D, 16L:8D and 24L:0D. At 21°C, the females increased spawning frequency, which was arrested at 11°C. Shorter light exposure at 21°C increased spawning frequency, while constant light inhibited and did not produce spawning. Constant dark (0L:24D) at 21°C produced the most (86%) spawns, but yielded poor larval quality. At 21°C with all photoperiod conditions except constant light, the first spawning took 94.8 ± 32.4 days to occur (n = 17). With females producing multiple spawns, the intervals between the first and second spawns and the second and third spawns were 37.7 ± 8.7 days (n = 6) and 31.0 ± 7.1 days (n = 2) respectively. Analysis of our data using response surface methodology (RSM) predicts the following conditions: at 15–19°C and 0–10 hr darkness for maximal survival and at 19–22°C and 0–8 hr darkness for spawning. The number of larvae produced was positively correlated with size (weight) of the female C. sapidus, suggesting the importance of female size in reproduction.  相似文献   

2.
Calanoid copepods, including species of the genus Acartia, are commonly used as larval diets for marine finfish. This study aimed to determine the separate effects of water temperature (18, 22, 24, 28° ± 0.5°C) and photoperiod (24L:0D; 18L:6D; 12L:12D; 8L:18D; 0L:24D) on Acartia grani egg production (EP), hatching rate (EHR) and population growth. Egg production rate was not affected by the two abiotic parameters. A. grani eggs incubated at T24°C and T28°C were the first to achieve 50% hatching rate (23–25 hr), with significant differences at the end of the experiment (48 hr) between T28°C treatment (EHR 88 ± 5%) and T18°C treatment (EHR 65 ± 2%). However, different temperature regimes did not affect final number of individuals in population growth experiment. Still, when eggs were excluded from data, population at lower temperatures (18°C) was mainly composed by the nauplii stage (72%), while at higher temperatures (24°C and 28°C) more than 60% of the population was composed by copepodites and adults. A. grani subjected to long‐day photoperiods had significantly lower EHR (16.7% at 24L:0D; 20.8% at 18L:6D) than at short‐day photoperiods (52.6% at 6L:18D; 50.0% at 0L:24D). In population growth experiment, eggs were the most common life stage after 12‐day culture. Lowest population number was found at constant light conditions (665.0 ± 197.1), suggesting higher metabolic rates and depletion of energy reserves in long‐day conditions. This study expanded knowledge on the biological response of A. grani to separate temperature and photoperiod regimes, and provided ground to improve the culture of this potential life feed species for hatcheries.  相似文献   

3.
The production of the flat oyster Ostrea edulis (L.) natural spat in Europe has decreased almost by 60% in the past ten years. Thus, the importance of the production of oyster spat in hatcheries is evident. One of the critical steps in hatchery production is broodstock conditioning, especially difficult in autumn, when gonadal development is in resting period. Conditioning is influence by temperature, photoperiod and nutrition. In this work, the effects of two temperature and three photoperiod regimes on the conditioning of O. edulis were studied for three years by stereological analyses and registering number and dates of spawning and larval yield. Temperature had a positive effect on the gonadal development of O. edulis during conditioning. The percentages of germinal cells in oysters conditioned with a gradient of temperature (14–18°C) were double compared to oysters conditioned at 15°C. Oysters conditioned with longer photoperiods showed higher percentages of germinal cells. There was no interaction between temperature and photoperiod. Spawning was observed in the oysters treated with daylight (8–16 h) ten weeks from the beginning of conditioning. Flat oysters conditioned with 8 h and 8–12 h of daylight delayed the first spawning for a month. Total larval production was higher in the oysters treated with the longest daylight gradient. Gonadal and gametogenic development was a non‐synchronic process and the spawning extended for around two months. A protocol for flat oyster broodstock conditioning in autumn by using both a gradient of temperature (14–18°C) and daylight (8–16 h) is proposed.  相似文献   

4.
Vitellogenesis is the process of production and uptake of vitellogenin (VtG), the precursor of vitellin (VT = yolk protein), for ovarian development. In Callinectes sapidus, hemolymph VtG levels are positively correlated with ovarian development. We aimed to develop an invasive tool to monitor ovarian development in order to predict spawning. The present study determined the effects of photoperiod and temperature conditions on levels of hemolymph VtG of the females in captivity kept in a closed recirculating system. Hemolymph VtG levels measured using a competitive ELISA correlates positively with temperature (21° versus 11°C) but are negatively correlated with photoperiod (0L:24D and 8L:16D versus 16L:8D and 24L:0D). Also, heavier (larger) females contain more VtG in their hemolymph and produce greater numbers of larvae. At 21°C, the VtG levels show high individual variations and continuously change over time, indicating that VtG production and uptake by ovary may be continuous and dynamic over multiple spawning events in C. sapidus. Microsatellite analysis of these females reveals a great diversity within and between the batches, implying that resident C. sapidus populations in the Chesapeake Bay are genetically dynamic. Such a great genetic diversity could be responsible for the high variation in VtG levels in these females.  相似文献   

5.
The effects of an artificially prolonged photoperiod on growth, survival rate, colouration, and sexual maturation in brook trout Salvelinus fontinalis during pre‐ and post‐spawning periods from 21 June to 06 November were investigated. Fish of mean initial weight ?150 g were reared at ambient photoperiod as well as with an artificially prolonged photoperiod produced by either a light‐emitting diode or a metal‐halide light. The fish groups subjected to a prolonged period of artificial light grew significantly larger and had a higher survival rate (p < 0.05), regardless of sex, and showed lower occurrence of fungal disease compared to controls reared in the natural photoperiod. We found a significantly higher number of sexually mature fish in the control groups compared with experimental groups. The increased photoperiod effectively delayed gonad development and increased somatic growth in both male and female brook trout, and also increased resistance to fungal disease. The increased photoperiod produced an observable difference in fish colouration, with control groups exhibiting more intensive spawning colouration.  相似文献   

6.
To investigate the effect of water temperature on gamete maturation and gamete quality, European grayling (Thymalus thymallus) were held under different temperature regimes prior to spawning. Maturation of males and females and their gamete quality depended strongly on temperature regime. The highest percentages of maturing fish and highest gamete quality were obtained under a creek water temperature regime with natural seasonal fluctuations. In warmed creek water (3–4°C), at a constant temperature of 8–9°C, and under an abruptly increasing temperature, regime maturation rates and gamete quality were reduced. The effect was more pronounced in females than in males. The spawning dates of different wild Austrian grayling populations were also correlated with water temperature data collected during the last 10–15 years. The mean spawning date expressed as the number of days from 21 December until spawning ranged from 98 to 111 days for lower elevation populations, while it was considerably delayed for an alpine population. All populations spawned at water temperatures of 5.5–7.2°C. Regression analysis indicated that spawning date correlated with (1) the overall mean water temperature from 21 December until spawning, (2) the mean water temperature during both the last 10 days and 15 days before spawning, and (3) the difference between mean January temperature and that of the last 15 days before spawning. The regression functions indicate that an increase in water temperature from 21 December to spawning of 1°C advances spawning by 5? days, and an increase of 1°C in the last 10–15 days advances spawning by 3? days.  相似文献   

7.
In order to clarify the respiratory responses strategy of Amur sturgeon Acipenser schrenckii exposed to water temperature changes, respiratory parameters of the fish were studied under two temperature regimes: fish acclimated at 13°C for Group I, temperature was increased to 16°C, 19°C, 22°C and 25°C and then returned stepwise to 22°C, 19°C, 16°C and 13°C; and fish acclimated at 25°C for Group II, the water temperature was reduced in steps to 22°C, 19°C, 16°C and 13°C, subsequently, returned to 16°C, 19°C, 22°C and 25°C. The results showed that the respiratory frequency (fR), oxygen consumption rate (VO2) and gill ventilation (VG) of the fish were directly dependent on the acute temperature in both acclimation groups (p < .05). The initial 25°C VO2 in Group II was significantly higher than the initial 13°C VO2 in Group I (p < .05), but was significantly lower than that at 25°C in Group I (p < .05). In Group I, respiratory stroke volume (VS.R) of fish significantly increased or decreased with the acute temperature increases or decreases, respectively (p < .05); oxygen consumption efficiencies (EO2) of fish did not significantly show differences when temperature increased to 25°C from 13°C (p > .05), but the EO2 significantly declined while returning to acclimation temperature (p < .05). In Group II, the VS.R of the fish did not significantly change with acute temperature fluctuations between 25 and 13°C (p > .05), while the EO2 increased with acute temperature increases (p < .05). The Q10 values for fR, VO2, VS.R, VG and EO2 were 1.53–1.72, 1.92–2.06, 1.07–1.60, 1.78–2.44 and 1.11–1.65 at 13–25°C of temperature interval respectively. Amur sturgeon showed partial metabolic compensation to temperature changes. The study results suggest that the ability of Amur sturgeon to regulate metabolism in response to acute temperature changes makes this species good adaptability in the aquaculture rearing.  相似文献   

8.
Adult red sea bream, Pagrus major (body weight, 1.0–2.0 kg) was exposed to three photoperiods [12 h light:12 h dark (12L:12D), 16 h light:8 h dark (16L:8D) and 24 h light:0 h dark (24L:0D)] from 2 months before spawning till the end of the spawning season to investigate growth, spawning and stress response. During the spawning season, tanks were checked every morning for spawned eggs. The growth performance in fish under 24L:0D was stimulated with significantly higher feed intake than those under other photoperiods (P<0.05). The number of eggs and gonadal histology confirmed that three and five females out of six in each of duplicate tanks of the 16L:8D treatment spawned. In contrast, only two out of six females in one tank of the 24L:0D treatment spawned, and no spawns were observed in the 12L:12D treatment. At the end of the spawning period, both 17β‐estradiol and testosterone levels were significantly higher in fish exposed to 16L:8D followed by 12L:12D and 24L:0D photoperiods (P<0.05). Photoperiod manipulation did not cause significant stress response in fish (P>0.05). The results suggest that stimulating the growth performance of red sea bream at reproductive stage with a 24L:0D photoperiod is possible if the fish are subjected to this photoperiod long before the onset of the spawning season.  相似文献   

9.
The influence of timing of extended photoperiods on growth and maturity of brook trout was investigated in a 112‐day experiment. The fish with mean initial weight of ~192 g were reared under four light regimes: one control group with natural ambient photoperiod and three groups exposed to an 18L:6D regime initiated at days 1, 23 or 46 of the growth trial. Light‐emitting diodes, with intensity of 250–1000 lux, depending on the distance from the light source, were used for extending light periods. There was a positive effect of prolonged day length on fish growth (< 0.05), and a delay in gonad development and sexual maturity. Significantly higher numbers of sexually mature fish were found among controls groups, regardless of sex. Survival rate was not affected by light regime. This study demonstrated that the short‐term expansion of the photo period delayed maturation and increased the growth rate of brook trout.  相似文献   

10.
The effect of extended photoperiods on growth and age at first maturity was investigated in 166 (79 females and 87 males) individually tagged Atlantic halibut Hippoglossus hippoglossus and in 114 (50 females and 64 males) individually tagged turbot Scophthalmus maximus. The halibut were reared at 11 °C on four different light regimes from 10 February to 6 July 1996: simulated natural photoperiod, (LDN), continuous light (LD24:0), constant 8 h light and 16 h darkness (LD8:16) and LD8:16 switched to continuous light 4 May 1996 (LD8:16–24:0). From 6 July 1996 to 9 February 1998 the LD24:0 and LD8:16–24:0 were reared together under continuous light and the LDN and LD8:16 together under natural photoperiod. The turbot were reared at 16 °C on three different light regimes: constant light (LD24:0), 16 h light:8 h darkness (LD16:8), or simulated natural photoperiod (LDN). After 6 months on the different photoperiods, the turbot was reared together on LDN for approximately 12 months until first maturation. Juveniles subjected to continuous light (halibut) or extended photoperiods (halibut and turbot) exhibited faster growth than those experiencing a natural photoperiod or a constant short day. Moreover, when the photoperiod increased naturally with day-length or when fish were abruptly switched from being reared on short-day conditions to continuous light, a subsequent increase in growth rate was observed. This growth enhancing effect of extended photoperiods was more apparent on a short time scale in Atlantic halibut than in turbot, but both species show significant long-term effects of extended photoperiods in the form of enhanced growth. In both species lower maturation of males was seen in groups exposed to extended or continuous light compared to LDN and this could be used to reduce precocious maturation in males leading to overall increase in somatic growth. This revised version was published online in August 2006 with corrections to the Cover Date.  相似文献   

11.
Farming of red tilapia is increasing rapidly. However, its commercial farming development is challenged by lack of clear information on genetic basis for skin colour and pigmentation differences due to environmental changes. This study investigated the effects of photoperiod (light:dark, L:D) on the growth and skin colour variation of Malaysian red tilapia. A total of 180 fish weighing 150.48 ± 0.44 g were reared under natural photoperiod (13L:11D, control), prolonged lightness (24L:0D) and prolonged darkness (0L:24D) in three replicates for 78 days. The weight gain of fish cultured under both prolonged light and darkness were significantly higher than fish under natural photoperiod. The tyrosinase level in ventral skin was significantly higher for fish cultured under prolonged darkness condition than in the other two photoperiod regimes. Contrary, the cysteine level in the dorsal skin was significantly higher in the fish cultured under natural photoperiod than in prolonged light and darkness. The relative mRNA expressions of SRY‐related HMG‐Box 10 (sox 10), tyrosine (tyr), tyrosine‐related protein 1 (tyrp‐1) and solute carrier family 7 member 11 (slc7a11) genes were significantly higher in ventral skin of fish under prolonged darkness than the other two photoperiods. This study demonstrates that photoperiod has an impact on melanogenesis and growth of red tilapia. Understanding the effects of photoperiod on genetic basis of red tilapia will help in selective breeding programme of the important economic traits for the development of commercial red tilapia farming.  相似文献   

12.
The combined effects of temperature (23 and 27 °C), light intensity (1100 and 2 lux or ≈ 21.5 and 0.04 μEinst m?2 s respectively) and photoperiod (10 h light/14 h dark and 14 h light/10 h dark) on ovarian maturation and spawning performance of ablated pond‐reared Penaeus merguiensis were investigated in a 51‐day experiment. The results showed that temperature was the most influential factor, followed by light intensity, whereas the effect of photoperiod was minor. As the maturation process progressed, the effects of temperature and light intensity became stronger while that of photoperiod became less significant. Temperature significantly affected all the reproductive parameters assessed. Prawns in the 27 °C treatments outperformed those in the 23 °C treatments (P < 0.05). The effect of light intensity was found to have a significant effect (P < 0.05) only on the proportion of spawners (prawns that spawned) and spawning rate. More prawns spawned at a higher rate under dim light (2 lux) than under strong light (1100 lux). Photoperiod affected only the proportion of prawns reaching stage III of ovarian development (P < 0.05). There were interactions between temperature and light intensity affecting the proportion of prawns reaching stage III and, although not significantly, the proportion of spawners (P = 0.177), fecundity (P = 0.134) and survival (P = 0.061). Overall, it is recommended that a combination of 27 °C, 2 lux and 10 h light/14 h dark is suitable for the maturation of pond‐reared P. merguiensis. There were indications that temperature can be used to control the rate of ovarian maturation. Also, it is possible to increase light intensity up to 1100 lux in P. merguiensis hatcheries if prawns are ablated.  相似文献   

13.
The purpose of this study was to investigate variations of glucose content, activities of enzymes involved in glycolysis and HSP70 in Litopenaeus vannamei subjected to one constant temperature (25°C) and four daily cyclical temperature change regimes (25 ± 1°C, 25 ± 2°C, 25 ± 3°C and 25 ± 4°C; max 12 am min 12 pm ). Both the glucose and HSP70 in treatment 25°C had a day/night rhythm city (L14:D10), but it gradually disappeared with the increase in temperature fluctuating amplitude. The PK activities varied more and more acutely with the increasing temperature fluctuating amplitude, especially, that in treatment 25 ± 4°C. HK activities were affected by the flux of glucose and the process of glycolysis, which tended to be stable with the increasing temperature fluctuating amplitude. Besides, the variations of PK activities were very abrupt at 25 ± 4°C, which might be unfavourable to the growth of shrimps. The temperature fluctuations affect metabolic adjustments and change the day/night rhythmicity of some physiological indicators.  相似文献   

14.
As a preliminary investigation into the effects of environmental factors on gonadal development using adaptive physiological characteristics, we examined the involvement of photoperiod and water temperature in the ovarian development of the red‐spotted grouper, Epinephelus akaara. During October 2014 (non‐breeding season), the photoperiod and water temperature were adjusted to 12L:12D and 18°C respectively. Every 3 weeks, daylight was increased as follows a 13L:11D and 14L:10D in the photo‐thermal treatment group, and the control group was maintained under natural conditions. After 9 weeks, water temperature was increased to 23°C in the photo‐thermal treatment group. In photo‐thermal treatment group, GSI increased significantly after 9 weeks (1.36 ± 0.19) and 12 weeks (3.31 ± 0.20) compared with the control group. At 12 weeks after photo‐thermal treatment, all fishes matured into ovaries containing late vitellogenic oocytes. The expression of FSHβ and LHβ mRNAs also increased significantly. However, in the control group, the ovarian development was maintained at an immature stage. Based on these results, we demonstrate that exposure to long photoperiod and high water temperature can induce the sexual maturation of immature red‐spotted grouper during the non‐breeding season.  相似文献   

15.
This study was conducted to evaluate the effects of photoperiod on the growth, feed conversion ratio (FCR) and spawning performance of Nile tilapia (Oreochromis niloticus) broodstock. Duplicate groups of two males (66±3.04 g) and six females (50.5±1.58 g) were stocked in 0.4 m3 fibreglass tanks in a recirculating water system at a male:female ratio of 1:3. The fish were subjected to four photoperiod treatments: 24:0 light:dark (L:D), 18L:6D, 12L:12D and 6L:18D. Light intensity was kept constant at about 2500 lx throughout the study. The fish were fed a commercial tilapia diet (35% crude protein and 16.6 MJ GE kg−1) at a daily rate of 2–3% of tank biomass, twice a day, for 130 days. Males grew significantly faster than females under all photoperiod regimes. The best growth rate and FCR were attained at 18L:6D, followed by 24L:0D, 12L:12D and 6L:18D respectively. The number of eggs per female, number of eggs per spawn and number of spawnings per female were all significantly higher in the 12L:12D treatment than in all other photoperiod cycles. Interspawning intervals and days elapsed per spawn were also shorter in the 12L:12D treatment. The time to first spawning was slightly longer in the 24L:0D and 6L:18D than in 12L:12D and 18L:6D light phases. The 18L:6D and 6L:18D photoperiods produced the lowest spawning performance. It is concluded that a 12L:12D photoperiod regime should be adopted for maximum fecundity, seed production and spawning frequencies of Nile tilapia broodstock reared in intensive, recirculating systems. If maximum reproduction is desired, a near‐natural day length photoperiod should be used.  相似文献   

16.
The present work investigates the importance of circadian and circannual rhythms in the photoperiodic control of reproduction in the rainbow trout. Maintenance of groups of 20–30 female trout under continuous light (LL), constant long (18L:6D) or short (6L:18D) days and conditions of constant temperature (8.5–9.0°C) and feeding rates (0.5% body weight, day–1), starting in February, produced markedly different spawning periodicities during the first and subsequent years of treatment. At the end of the first year, spawning was advanced by up to 2 months in the fish under LL and 18L:6D and delayed by up to 5 months in those under 6L:18D when compared with the December spawning of control fish under ambient light. Continued exposure of the fish to the same photoperiod regimes produced cycles of spawning and peak levels of testosterone, oestradiol-17 and calcium (as an index of vitellogenin) at intervals of 150–170 days under LL and 18L:6D and 320–420 days under 6L:18D. The functional importance of these spawning cycles and their relationship to circannual rhythms and the control of reproduction is discussed.Exposure of fish to skeleton (6L:4D:2L:12D, 6L:6D:2L:10D and 6L:8D:2L:8D) and resonance (6L:42D, 6L:48D and 6L:54D) procedures produced ranges of spawning times up to two months in advance of control fish. Results with the resonance regimes, where fish received only a half the light-dark cycles and a quarter of the total daylight hours of those on ambient light cycles show that trout do not measure daylength and time by counting daily cycles or by hour-glass mechanisms. Collectively, these data support the proposition that rhythmic processes of photosensitivity are involved in the modulation of reproductive development in the rainbow trout and that circadian and circannual rhythms cooperate in the timing and entrainment of this cycle.  相似文献   

17.
A major problem in tilapia aquaculture is the frequent reproduction of female fish, leading to increased competition for supplemented food and stunted somatic growth. The feasibility of using photoperiod manipulation to arrest the reproductive performance of tilapia Oreochromis niloticus was therefore examined. Newly hatched O. niloticus were reared in the laboratory under 12L:12D photoperiod at 28 °C. Fish (230–340 g) were maintained under 6L:6D, 12L:12D and 14L:10D photoperiod at 28 °C during the course of this study. Effect of photoperiod manipulation on reproductive parameters of fecundity, gamete quality, offspring viability and overall reproductive success were evaluated. Steroid levels (estradiol-17β, E2; testosterone, T) during the spawning cycles of fish were analyzed on days 1 and 3 postspawning and at 3-day intervals thereafter. A total of 72 female fish from each photoperiod treatment were investigated the changing pattern of E2, T and GSI with fish that have spawned once, twice and three times. Fish exposed to 12L:12D and 14L:10D photoperiod spawned successfully throughout the study. Whereas the spawning of fish exposed to 6L:6D photoperiod was arrested after three to four spawning cycles. The arrestment of spawning in fish exposed to 6L:6D photoperiod was paralleled by a significant decrease in plasma levels of E2 (P<0.05). By contrast, there was no major difference in T levels among the treatments. These findings suggest that photoperiod manipulation can be used to arrest the spawning in tilapia O. niloticus. The utility of this in controlling the problem of overcrowding due to excess offspring in tilapia aquaculture is applicable.  相似文献   

18.
Shi drum (Umbrina cirrosa L.) larvae were maintained under three photoperiod regimes: a natural photoperiod regime (16L:8D), continuous light (24L) and equal durations of light and dark (12L:12D) from the end of the vitelline phase to the end of the metamorphosis. Muscle and body parameters were studied at hatching and at 4, 10, 14, 39 and 55 days post hatching (dph). During the vitelline phase, the total body length growth was scarce, whereas the muscle grew significantly, being the hypertrophy of the main mechanism involved. Both the total body length and the hypertrophy were significantly greater at 16L:8D than in the rest of photoperiod regimes. At 10 and 14 dph, the greatest body length was reached at 16L:8D, followed by the 24L group, showing the 12L:12D group the lowest values. At 14 dph, the hypertrophy and hyperplasia were also higher at 16L:8D than in the rest of groups. At 39 dph, the highest values of body length were reached in both 16L:8D and 12L:12D regimes, this latter group reaching the highest values of hypertrophy, thus showing a compensatory growth when comparing with the previous stages. The end of the metamorphosis took place at 50–55 dph in all the groups, with 2.7–3.1 cm of body length (> 0.05). At this stage, the transverse area of the white muscle was similar among the groups, but the greatest hypertrophy was reached at 16L:8D, whereas the highest hyperplasia was reached at 24L.  相似文献   

19.
20.
In Channa punctatus maintained under ambient photothermal conditions (10.5L:13.5D; 13 ± 2°C) in the postspawning season, hypothalamic dopamine (DA) and norepinephrine (NE) content varied diurnally, but serotonin (5-HT) and monoamine oxidase (MAO) did not. Exposure of the fish to high temperature (25 ± 2°C), with or without altering the photopheriod, increased the NE and DA content, and NE turnover but decreased DA turnover. Increasing the photoperiod (16L:8D) alone had no significant effect on catecholaminergic (CA) content or activity. When the fish were exposed to long photoperiod, with or without altering the temperature, a day-night variation in 5-HT content and activity was induced in the postspawning season. An increase in temperature alone had no significant effect on 5-HT content or activity. Hypothalamic MAO activity was elevated in fish exposed to high temperature alone, or in combination with long photoperiod, but was not affected significantly by long photoperiod treatment. These results indicate that CA activity is influenced largely by ambient high temperature, whereas serotonergic activity is controlled primarily by photoperiod, with high temperature having an additive effect. MAO activity seems to be influenced by both temperature and photoperiod in this species.  相似文献   

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