J. Appl. Ichthyol.

23 (2007), 621623 2007 The Authors Journal compilation 2007 Blackwell Verlag, Berlin ISSN 01758659

Received: March 14, 2006 Accepted: November 1, 2006 doi: 10.1111/j.1439-0426.2007.00856.x

Short communication The eect of photoperiod on the reproduction of Chirostoma estor estor Jordan 1879 tzcuaro, Mexico from Lago de Pa
vez-Sosa1, V. O. Santoyo-Guzma n1, A. Campos-Mendoza1, C. C. MartinezBy C. A. Mart nez-Palacios1, J. C. Cha 2 2 Chavez and L. G. Ross
Laboratorio de Acuicultura y Nutricion, INIRENA, Universidad Michoacana de San Nicolas de Hidalgo, Morelia, Michoacan, Mexico; 2Institute of Aquaculture, University of Stirling, Stirling, UK
1

Introduction The Mexican silverside (Chirostoma estor estor) is one of a group of native Atherinopsid freshwater species endemic to the lakes of Central Mexico. C. estor estor is the principal tzcuaro where for many years species of the shery in Lake Pa it has provided a subsistence income for a large number of families. It is a high-value species that has become endangered because of pollution, introduction of exotic species and through over-shing (Mart nez-Palacios et al., 2002, 2004). It has great potential for aquaculture and in recent years there has been increased interest in its culture. The development of native species requires comprehensive control of their reproductive cycle. C. estor estor is able to spawn throughout the year given the proper circumstances, but its reproductive peak in the wild is from February to July. In many species it is very advantageous to be able to manipulate the reproductive activity of broodstock to improve egg and sperm production (Bromage et al., 2001). This may enable continuous yearround production of fry without dependence upon the natural environment. In recent years photoperiod manipulation in sh has been shown to result in better growth, early maturation and yearround production in several annual and temperate species (Bromage et al., 2001). Studies on salmon with additional light have been shown to enhance growth in sea cages (Endal et al., 2000), while exposure to short winter day-lengths helps trigger precocious maturation and smoltication (Berrill et al., 2003). This is also true for male masu salmon (Oncorhynchus masou) in which testicular maturation is accelerated by a short photoperiod (8L : 16D) and delayed by long photoperiods (16L : 8D) (Amano et al., 2000). In rainbow trout, Bromage et al. (1984) and Scott et al. (1984) found that constant long days or continuous light both advanced spawning, and Davies et al. (1999) found that a combination of long and short photoperiods (18L : 6D and then 6L : 18D) can advance or delay maturation and spawning. It is well accepted that photoperiod aects reproduction in sh in one way or another; it is considered the most important environmental cue for breeding seasonality and maturation. Recent work on multiple batch spawning Nile tilapia (O. niloticus L.), has shown that long photoperiods (i.e. 18L : 6D) promote general spawning performance (total/relative fecundity, ISI) in female broodstock compared with short (6L : 18D) and normal (12L : 12D) days (Campos-Mendoza et al., 2004). It has also been observed that a 6L : 6D
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photoperiod has a positive eect on sh growth but is detrimental to egg quality with no viable eggs being produced after three cycles under this regime (Biswas and Takeuchi, 2003; Biswas et al., 2004, 2005). Photoperiod also seems to aect earlier stages of tilapia as shown by El-Sayed and Kawanna (2004) when fry but not ngerlings were signicantly aected by longer days (i.e. 18L : 6D and 24L). A similar paper (Rad et al., 2006) showed a signicant dierence only during the ngerling stage. Fish are very sensitive to light, and although thresholds needed for reproduction (maturation) are not yet known (Bromage et al., 2001), in general it has been found that high intensity light levels are required for growth optimization (Boeuf and Le Bail, 1999) while shorter or longer photoperiods are needed to stimulate maturation in sh (Amano et al., 2000; Bromage et al., 2001; Berrill et al., 2003). Materials and methods Since 2003, C. estor estor broodstock have bred successfully in 4 m diameter tanks in our laboratory, providing a source of juveniles completely independent from the wild stocks. From eld observations made during shing trips to capture broodstock it became clear that the lunar cycle had a marked inuence on reproductive behaviour. During the full moon more females migrated to the lake shore to spawn and those females captured by the shermen contained a greater number of mature eggs. Lake Patzcuaro is located at 1937 latitude (North) and 10137 longitude (West); at these latitudes, there is relatively little variation in photoperiod, with the longest days (14L : 10D) (L: light: D: dark) occurring during summer and the shortest days (10L : 14D) in winter. Although the number of daylight hours appears to be fairly constant during the year, moon cycles and other environmental cues such as temperature could have a major inuence on the species seasonality. Based upon this and our observations with the species, an experiment was designed to investigate the eect of day length on reproductive output in C. estor estor. Light-proof chambers were constructed with four dierent photoperiods: 6L : 18D, (L, light; D, dark), 12L : 12D, 18L : 6D and 24L : 0D. Each treatment consisted of 12 males and 12 females (mean weight 100 g). The experiment was continued for 7 months between December 2003 and June 2004, with egg production monitored daily. After observing reproductive output in these photopewww.blackwell-synergy.com

01758659/2007/23050621$15.00/0

622 Table 1 Experimental conditions and water quality parameters in two Chirostoma estor estor trials Parameter Trial dates Light regime (dark : night) Light intensity at surface (lux) Tank dimensions (length width height) (m) Tank volume (m3) Flow rate (l min)1) Stocking level Mean initial body weight (g) Temperature (C) pH Dissolved oxygen (mg l)1)2
1 2

C. A. Mart nez-Palacios et al.

Natural photoperiod trial May 2004October 2004 Natural photoperiod1 Natural daylight 2.50 1.90 0.90 4.2 12 10# : 20$ 100 24 1 9 0.1 >6
1

Controlled photoperiod trial December 2003June 2004 6D : 18L 75 2.10 1.50 0.90 2.8 12 12# : 12$ 100 22 1 9 0.1 >6 12D : 12L 75 2.10 1.50 0.90 2.8 12 12# : 12$ 100 22 1 9 0.1 >6 18D : 6L 75 2.10 1.50 0.90 2.8 12 12# : 12$ 100 22 1 9 0.1 >6 24D : 6L 75 2.10 1.50 0.90 2.8 12 12# : 12$ 100 22 1 9 0.1 >6

Supplemented by low-level background lighting at night. Altitude 1900 m.

riods for 5 months, all sh groups were re-established in a photoperiod of 18L : 6D. A further experiment was designed to investigate the natural reproductive behaviour of 30 adult C. estor estor, maintained at a sex ratio of 2 : 1 female : male, in two 2.8 m3 concrete tanks. Reproductive output was monitored for 6 months, from May to October 2004. Experimental conditions in both experiments are summarized in Table 1. Results and discussion In the photoperiod trial the highest fecundity and number of spawnings were obtained with the long day length treatments (24L : 0D and 18L : 6D). In the short day length treatments (12L : 12D and 6L : 18D), no spawnings were obtained throughout the trial apart from one very small spawning at the beginning of the experiment at 12L : 12D (Fig. 1). It was very dicult to elucidate the number of eggs spawned by a single female, as the sh were stocked together in each trial. The species is a multiple-spawner and to identify the number of eggs produced by a particular female they would need to be stocked individually. After changing to the 18L : 6D photoperiod all groups spawned successfully. The change was most marked in the short day length groups which had not spawned

at all during the ve previous months. This observation showed that the change to longer day length stimulated eciently the reproductive activity of C. estor estor. In the second trial there was no discernible relationship between lunar cycle and egg production (Fig. 2), with eggs being produced in both the light and dark phases of the lunar month. It was notable, however, that eggs produced from July onward were out of season; this may have been due to an extended day produced by high background lighting adjacent to the tanks. This result suggested that during the reproductive season of this species, sh may spawn during extended light periods (long days) caused by augmentation of the solar day during full moon periods. The length of daily illumination may act as a trigger for spawning, possibly by acting on hormonal rhythms (Bromage et al., 2001). Extended days may be the cue to spawning in C. estor estor, as even in the winter months the species will spawn at very low temperatures (1214C). Conclusion In summary, these preliminary experiments have shown that light is a major factor in the control of reproductive activity of C. estor estor, oering a means of managing egg production. Recently, similar results have also been obtained in the laboratory with the closely related species Chirostoma promelas

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Fig. 1. Total fecundity in Chirostoma estor estor pre-conditioned to dierent experimental photoperiods and following a change to 18 h light

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NM = New moon, FQ = First quarter, FM = Full moon, LS = Last quarter

Fig. 2. Number of Chirostoma estor estor eggs per spawning per month and by lunar phase

mb er Se pte mb LQ er Se pte mb NM er Se pte mb er FQ Oc tob er FM

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Chirostoma photoperiod and reproduction

623 Bromage, N. R.; Porter, M.; Randall, C., 2001: The environmental regulation of maturation in farmed nsh with special reference to the role of photoperiod and melatonin. Aquaculture 197, 6398. Campos-Mendoza, A.; McAndrew, B. J.; Coward, K.; Bromage, N., 2004: Reproductive response of Nile tilapia (Oreochromis niloticus) to photoperiodic manipulation; eects on spawning periodicity, fecundity and egg size*1. Aquaculture 231, 299314. Davies, B.; Bromage, N.; Swanson, P., 1999: The brainpituitary gonadal axis of female rainbow trout Oncorhynchus mykiss: eects of photoperiod manipulation. Gen. Comp. Endocrinol. 115, 155 166. El-Sayed, A.-F. M.; Kawanna, M., 2004: Eects of photoperiod on the performance of farmed Nile tilapia Oreochromis niloticus: I. Growth, feed utilization eciency and survival of fry and ngerlings. Aquaculture 231, 393402. Endal, H. P.; Taranger, G. L.; Stefansson, S. O.; Hansen, T., 2000: Eects of continuous additional light on growth and sexual maturity in Atlantic salmon, Salmo salar, reared in sea cages. Aquaculture 191, 337349. Mart nez-Palacios, C. A.; Barriga Tovar, E.; Taylor, J. F.; Rios Duran, G.; Ross, L. G., 2002: Eect of temperature on growth and survival of Chirostoma estor estor Jordan 1879, monitored using a simple video technique from remote measurement of length and mass of larval and juvenile shes. Aquaculture 209, 369377. Mart nez-Palacios, C. A.; Comes-Morte, J.; Tello-Ballinas, J. A.; Toledo-Cuevas, M.; Ross, L. G., 2004: The eects of saline environments on survival and growth of eggs and larvae of Chirostoma estor estor Jordan 1879. (Pisces: Atherinidae). Aquaculture 238, 509522. Rad, F.; Bozaoglu, S.; Ergene Gozukara, S.; Karahan, A.; Kurt, G., 2006. Eects of dierent long-day photoperiods on somatic growth and gonadal development in Nile tilapia (Oreochromis niloticus L.). Aquaculture 255, 292300. Scott, A. P.; Baynes, S. M.; Skarphedinsson, O.; Bye, V. J., 1984: Control of spawning time in rainbow trout, Salmo gairdneri, using constant long daylengths. Aquaculture 43, 225233. Authors address: Carlos A. Mart nez-Palacios, Laboratorio de Acuicultura y Nutricion, INIRENA, Universidad s de Hidalgo, Morelia, Michoacana de San Nicola n, Me xico. Michoaca E-mail: fobospalacios@hotmail.com

Jordan and Snyder 1899. Clearly, more information is needed to fully understand the eect of photoperiod on egg quality as well as the relationship between photoperiod, cycles of melatonin production and release and the involvement of the HPI axis. The use of photoperiod to manipulate spawning is already being used to provide a year-round supply of C. estor estor. This will make a major contribution to development of aquaculture of the species without the necessity of relying upon natural, wild spawning. Acknowledgement We wish to thank CONACyT, Mexico and DEFRA, UK, for nancial support. References
Amano, M.; Iigo, M.; Ikuta, K.; Kitamura, S.; Yamada, H.; Yamamori, K., 2000: Roles of melatonin in gonadal maturation of underyearling precocious male masu salmon. Gen. Comp. Endocr. 120, 190197. Berrill, I. K.; Porter, M. J. R.; Smart, A.; Mitchell, D.; Bromage, N. R., 2003: Photoperiodic eects on precocious maturation, growth and smoltication in Atlantic salmon, Salmo salar. Aquaculture 222, 239252. Biswas, A. K.; Takeuchi, T., 2003: Eects of photoperiod and feeding interval on food intake and growth rate of Nile tilapia Oreochromis niloticus L. Fish. Sci. 69, 10101016. Biswas, A. K.; Maita, M.; Yoshizaki, G.; Takeuchi, T., 2004: Physiological responses in Nile tilapia exposed to dierent photoperiod regimes. J. Fish Biol. 65, 811821. Biswas, A. K.; Morita, T.; Yoshizaki, G.; Maita, M.; Takeuchi, T., 2005: Control of reproduction in Nile tilapia Oreochromis niloticus (L.) by photoperiod manipulation. Aquaculture 243, 229239. Boeuf, G.; Le Bail, P. Y., 1999: Does light have an inuence on sh growth? Aquaculture 177, 129152. Bromage, N. R.; Elliott, J. A. K.; Springate, J. R. C.; Whitehead, C., 1984: The eects of constant photoperiods on the timing of spawning in the rainbow trout. Aquaculture 43, 213223.