Printer Friendly

Variation in Advertisement Call Properties of the Japanese Quail.

Byline: M. Sezer and O. Tekelioglu

Abstract

The aim of this study was to investigate the amount variation in four spectral and six temporal properties of advertisement calls between and within- male Japanese quail and to establish the effect of age of sexual maturation to these properties. The first, second, third syllables and damping parts were 68.2, 74.3, 501.1 and 95.9 ms long, on average, and had peak frequencies of 1719.9, 1989.9, 2203.2 and 2352.7 Hz, respectively. There was high between individual variations in all call traits. In relation to within-male variation, there were high medians and low ranges for spectral properties while low medians and high ranges for temporal properties. Moderate positive correlations between syllable lengths and age of sexual maturation, low negative correlations between interval lengths and age of sexual maturation were observed.

Repeatability of the advertisement call characteristics were high, but estimates for spectral properties were slightly lower (0.66-0.82) than that for temporal properties (0 89-0.96). These results suggest that temporal call parameters have the potential to be used in individual identity and these traits might carry information used by females in mate choice. Spectral properties could be recognized as stable call properties that may be used in efficient transmission or species recognition.

Keywords: Quail, crow, individual identity, call parameters, repeatability.

INTRODUCTION

When visual communication is limited, birds mainly use the acoustic channel to communicate and a few features of the song that are resistant to degradation are generally used for species-specific recognition (Aubin et al., 2004). This coding system allows the bird to transmit specific information at long ranges. Between-male variation in secondary sexual traits influences success in sexual competition to gain mating opportunities with choosy females (Andersson, 1994). Hence, examination of both between and within-individual variation is required to understand the evolution of sexual behaviours (Boake, 1994). One way of quantifying between-individual variation in a phenotypic trait while taking within-individual variation into account is to use the quantitative genetic concept of repeatability (Falconer and Mackay, 1981; Boake, 1989). Higher variation between individuals compared to that within individuals, i.e. high repeatability, suggests the possibility of high heritability values (Boake, 1989).

Traits with important signalling functions in mate choice are expected to show higher repeatability than less-critical traits. The Japanese quail was first domesticated in the 11th century as a songbird for advertisement calls of males and still lives in the wild in Asia (Cheng and Kimura, 1990). The Japanese quail was recognized as a food source at the beginning of the 20th century and farmed for commercial meat and egg production (Mills et al., 1997). Besides its own merit in fancy and commercial breeding, Japanese quail is a valuable model system for a variety of scientific studies. Male Japanese quails produce territorial songs that consist of three syllables; two short trills followed by an extended syllable that fades away with a short damping component. Guyomarc'h et al. (1998) reported that the temporal patterns of these calls were highly stereotyped and could play a role in inter- individual recognition.

The role of male mating calls in female mate choice and indication of male advertisement calls containing species-specific and individual information about the caller were also reported (Guyomarc'h et al., 1998; Deregnaucourt et al. , 2001; Deregnaucourt and Guyomarc'h, 2003). On the other hand, age and reproductive status of the males are the factors affecting the syllable and interval lengths of the calls (Schleidt and Shalter, 1973; Deregnaucourt et al., 2009). There is no influence of learning on Galliforms' call and it is believed that their call features are mainly determined by the genetic constitution of the individual (Deregnaucourt et al., 2001).

The aims of this study were two fold. The first objective was to investigate the amount of variation in advertisement call properties between and within male Japanese quail to estimate the repeatability because both sources of trait variation can influence the course of sexual selection. The second aim was to test the age of sexual maturity effect on the call properties.

MATERIALS AND METHODS

The data for this study were obtained from the Japanese quail population (Coturnix coturnix japonica) at the Quail Breeding Unit of Gaziosmanpasa University, Tokat, Turkey. Birds used in this study were obtained from five consecutive hatchings by artificial incubation. Analyses were carried out with the records of 1514 advertisement calls obtained from randomly selected 393 male Japanese quails from the study population. We kept the birds in mixed gender groups of 15 to 20 individuals to allow the males in contact with females and have physical interactions with other individuals. Chicks were housed for the first three weeks under 24 hour light, with following weeks at 16:8 light:dark cycle. The temperature started at 36degC. Temperature was decreased by 3degC every week until it reached 24degC. Birds were allowed ad libitum access to food and water.

They were fed with 240 g/kg crude protein (CP) and 13.39 MJ ME/kg starter diet for 21 days, 190 g/kg CP and 12.55 MJ ME/kg grower diet between 22 and 35 days of age and thereafter 170 g/kg CP and 11.50 MJ ME/kg breeder diet.

Recordings

The foam produced by the cloacal gland is an androgen dependent secondary sexual character in the male Japanese quail and could be used as an external index of sexual maturity (Mohan et al., 2002; Marin and Satterlee, 2004). Males were inspected daily (after 25 days of age), to determine the day of cloacal gland foam production, and this day was recorded as the age of sexual maturity (ASM). Calls of males used in this study were recorded when the individuals were 8 week of age and they were completely sexually mature, as testified by the development of the proctodeal gland with foam production. Males were placed in a sound-proof chamber (1x1.5x0.5m) for a period of five minutes and their crows were recorded. When recording of a male call was unsuccessful, the procedure was repeated two days after the first attempt. All males were recorded within 2 weeks. Advertisement calls were recorded with a personal computer (sampling rate=32 kHz) equipped with a Sennheiser MD 41 dynamic microphone.

Fig. 1. Oscillogram of a male Japanese quail call. S1, Int1, S2, Int2, S3 and Dm are the lengths of the first syllable, interval one, the second syllable, interval two, the third syllable and damping, respectively.

Acoustic analyses

Japanese quail's calls are composed of three parts; two short syllables followed by an extended trill that ends with a short component that was called 'damping' (Fig. 1). The damping (Dm) is a measure of the fading of vibratory motion of the third syllable. Hence, each recorded song was analyzed to measure six temporal [durations of syllables (S1, S2 and S3), intervals (Int1, Int2) and damping (Dm)] and four spectral properties [peak frequencies of syllables (Fr1, Fr2, Fr3) and damping (FrD) component]. The peak frequency displays the frequency of the strongest spectral component in the selected entire syllable.

Statistical analyses

Variability in each call property between and within males was expressed as coefficients of variation. Coefficients of variation between males were based on average values for each male. The level of association between each pair of call variables and ASM was determined by correlation analyses. Only males for which we had at least three crows were included in ANOVA analyses to estimate the repeatability of male song properties. Corrected sample size (n0), that is a coefficient related to the sample size per group in the analysis of variance, and standard errors for the repeatability estimates were calculated following the procedures outlined by Lessells and Boag (1987) and Becker (1992).

RESULTS

The statistical description and variability of call properties is summarized in Table 1. Sample sizes for the examined traits vary because all parameters could not be measured for every call. For example, relatively small data size in Fr1 component mainly arose from the lack of first syllable of some males. The length of the first component (Int1) was over two times longer than the length of the second interval (Int2). The first and second syllable length was short and near to each other but the third syllable was remarkably long. Interestingly, component frequencies increased consistently throughout the call. Variation expressed in terms of the coefficient of variation (Table I), was higher for the lengths of syllables and intervals than for spectral properties of the syllables (Mann-Whitney test: U=0.0, Z = -2.558, P 0.05). This could be expected because regardless of the ASM all males were recorded within 8 to 10 weeks of age.

Temporal properties of the song tabulated within sexual maturation groups were presented in Figures 2 and 3. Because, the effect of ASM on spectral

Table I.- Between and within-male variability in calling song properties.###

Property###Between male###CV (%) within male

###N###Mean###SD###Min###Max###CV(%)###Median###Range

Int1###336###56.0###29.03###5.2###229.5###51.82###11.78###23.83

Int2###373###113.5###43.63###10.3###322.3###38.42###6.79###24.75

S1###336###68.2###20.17###29.2###150.0###29.56###7.93###24.01

S2###372###74.3###29.94###22.8###230.5###40.31###9.81###32.89

S3###393###501.1###143.13###211.3###1109.5###28.56###4.30###13.49

Dm###393###95.9###49.38###17.4###265.0###51.50###13.07###42.43

Fr1###311###1719.9###209.39###842.2###2712.9###12.17###4.66###24.34

Fr2###350###1989.9###324.59###1125.5###3103.3###16.31###6.72###32.31

Fr3###387###2203.2###382.12###1568.5###4164.0###17.34###4.72###35.78

FrD###365###2352.7###496.82###1175.8###3676.0###21.12###4.64###39.53

S1, S2 and S3 are the duration of the first, second and third syllable respectively; Int1 and Int2 are the durations of interval one and two; Dm is the damping duration. Fr1, Fr2, Fr3 and FrD are the peak frequencies of the first, second, third syllables and damping part, respectively. Scale for duration and frequency measurements are in "ms" and "Hz", respectively.

Table II.- Variation and repeatability (r) of male call properties.

Properties###Mean Square###Fa###N###n0###r###SE###

###Between###Within###

In1###50.54###3097.56###61.29###336###3.85###0.94###0.025###

In2###83.17###7048.86###84.75###373###3.83###0.96###0.019###

S1###45.48###1515.30###33.31###336###3.85###0.89###0.043###

S2###77.76###3376.47###43.42###372###3.83###0.92###0.034###

S3###833.38###77826.62###93.39###393###3.83###0.96###0.017###

Dm###233.77###9301.89###39.79###393###3.83###0.91###0.037###

Fr1###14063.37###164906.36###11.73###311###3.84###0.74###0.093###

Fr2###45550.95###388285.20###8.52###350###3.81###0.66###0.111###

Fr3###36322.56###549623.69###15.13###387###3.75###0.79###0.078###

FrD###51584.71###938364.50###18.19###365###3.81###0.82###0.069

a for all properties P<0.001

All of the repeatability estimates for the advertisement call characteristics were high (Table II), but estimates for spectral properties (frequencies of the components) were slightly lower (Mann-Whitney test: U=0.0, Z = -2.566, P < 0.05) than that for temporal properties (intervals and syllables lengths).

DISCUSSION

Our results indicate that temporal call parameters have the potential to be used in individual identity. On the other hand, spectral properties had low between individuals variation hence they could be recognized as more stable properties of the call. This could reflect a history of stabilising selection on advertisement calls for efficient transmission or species recognition.

In our study, all mean values of the temporal call properties were lower than reported by Collins and Goldsmith (1998). These differences could be the result of genetic differences between the populations. We examined the third syllable of the call excluding the damping component. When the damping duration was added to the duration of the third syllable, it was still 20% lower than their report. On the other hand, the values for peak frequencies of the syllables were higher than those reported by Collins and Goldsmith (1998). Variation in the advertisement calls of animals and the potential of vocal parameters to be used in individual recognition has been proposed by many authors (Forrest and Green, 1991; Mathevon, 1996; Collins and Goldsmith, 1998; Mathevon et al., 2003; Sharp and Hatchwell, 2005). In this study, the pattern of between-male CVs indicated different repeatability patterns for the spectral and temporal properties of the call.

The high repeatability of song properties of male Japanese quail was due to increased variation between males rather than to decreased variation within individuals. Hence, estimated high repeatability for call properties may not indicate a correlated high heritability, but moderate heritability estimates might be expected.

The call properties that are more variable between-individuals than within-individuals can potentially be used in sexual selection as individual markers (Boake, 1994; Johnsen and Zuk, 1996; Hack, 1998; Aubin el al., 2004). It has also been reported that female Japanese quail express a preference among males (Deregnaucourt and Guyomarc'h, 2003; Ophir and Galef, 2003). Calls of male Japanese quail may be used by females to select among males, especially if they reflect aspects of male quality. Although repeatability of the frequency of the third syllable (0.19) and the duration of the first syllable (0.04) were unexpectedly low, Collins and Goldsmith (1998) reported high repeatability estimates for the vocal parameters of the Japanese quails.

Age and reproductive status of the males have been reported as factors affecting the temporal properties of the quail calls (Schleidt and Shalter, 1973; Deregnaucourt et al., 2009) . It is also known that males with genetically high growth potential reach the sexual maturity faster than the ones with low growth potential (Sezer et al., 2006). Hence, this preliminary estimate of repeatability indicated that upcoming genetic analysis of the Japanese quail song properties would be feasible when they are based on recordings which take into account the age of sexual maturation of the examined males. Males used in this study were recorded within a five-minute at the very beginning of their sexual maturation. It would be interesting to measure repeatability over long time scales to test the consistency of a male's calls across years or seasons in future studies.

Additional studies exploiting call properties that are important for individual recognition and their genetic basis are also required to understand the selection pressure on quail calls.

REFERENCES

ANDERSSON, M., 1994. Sexual selection. Princeton, New Jersey.

AUBIN, T., MATHEVON, N., DA SILVA, M.L., VIELLIARD, J.M.E. AND SEBE, F., 2004. How a simple and stereotyped acoustic signal transmits individual information: the song of the White-browed Warbler Basileuterus leucoblepharus. Anais Acad. Brasil. Cienc., 76: 335-344.

BECKER, W.A., 1992. Manual of quantitative genetics. WA Academic Enterprises, Pullman.

BOAKE, C.R.B., 1989. Repeatability: its role in evolutionary studies of mating behavior. Evolut. Ecol., 3: 173-182.

BOAKE, C.R.B., 1994. Behavioral variation and speciation: flexibility may be a constraint. In: Flexibility and constraint in behavioral systems (eds. R.J. Greenspan and C.P. Kyriacou), pp. 259-268. John Wiley, New York.

CHENG, K.M. AND KIMURA, M., 1990. Mutations and major variants in Japanese quail. In: Poultry breeding and genetics (ed. R.D. Crawford), pp. 333-362, Elsevier, New York.

COLINS, S.A. AND GOLDSMITH, A.R., 1998. Individual and species differences in quail calls (Coturnix c. Japonica, C. c coturnix and a hybird). Ethology, 104: 977-990.

DEREGNAUCOURT, S. AND GUYOMARC'H, J.C., 2003. Mating Call Discrimination in Female European (Coturnix c. coturnix) and Japanese Quail (Coturnix c. japonica). Ethology, 109: 107-119.

DEREGNAUCOURT, S., GUYOMARC'H, J.C. AND RICHARD, V., 2001. Classification of hybrid crows in quail using artificial neural networks. Behav. Proc., 56: 103-112.

DEREGNAUCOURT, S., SAAR, S. AND GAHR, M., 2009. 586 M. SEZER AND O. TEKELIOGLU Dynamics of crowing development in the domestic Japanese quail (Coturnix coturnix japonica). Proc. biol. Sci., 276: 2153-2162.

FALCONER, D.S. AND MACKAY, T.F.C., 1996. Introduction to quantitative genetics, 4th edn. Longman: London.

FORREST, T.G. AND GREEN, D.M., 1991. Sexual selection and female choice in mole crickets (Scapteriscus: Gryllotalpidae): Modelling the effects of intensity and male spacing. Bioacoustics, 3: 93-109.

GUYOMARC'H, J.C., AUPIAIS, A. AND GUYOMARC'H, C., 1998. Individual differences in the long-distance vocalizations used during pair bonding in European quail (Coturnix coturnix). Ethol. Ecol. Evolut., 10: 333-346.

HACK, M.A., 1998. The energetics of male mating strategies in field crickets (Orthoptera: Gryllinae: Gryllidae). J. Insect Behav., 11: 853-867.

JOHNSEN, T.S. AND ZUK, M., 1996. Repeatability of mate choice in female red jungle fowl. Behav. Ecol., 7: 243-246.

LESSELLS, C.M. AND BOAG, P.T., 1987. Unrepeatable repeatabilities: a common mistake. The Auk., 104: 116-121.

MARIN, R.H. AND SATTERLEE, D.G., 2004. Cloacal gland and testes development in male Japanese quail selected for divergent adrenocortical responsiveness. Poult. Sci., 83: 1028-1034.

MATHEVON, N., 1996. What parameters can be used for individual acoustic recognition by the Greater Flamingo? Compt. rend. Acad. Sci. Serie III, Sciences de la vie. 319: 29-32.

MATHEVON, N., CHARRIER, I. AND JOUVENTIN, P., 2003. Potential for individual recognition in acoustic signals: a comparative study of two gulls with different nesting patterns Ethology, 326: 329-337.

MILLS, A.D., CRAWFORD, L.L., DOMJAN, M. AND FAURE, J.M., 1997. The behavior of the Japanese or domestic quail (Coturnix japonica). Neurosci. Biobehav. Rev., 21: 261-281.

MOHAN, J., MOUDGAL, R.P., VENKATA, K., SASTRY, H., TYAGI, J. AND SINGH, R., 2002. Effects of hemicastration and castration on foam production and its relationship with fertility in male Japanese quail. Theriogenology, 58: 29-39.

OPHIR, A.G. AND GALEF, B.G., 2003. Female Japanese quail that 'eavesdrop' on fighting males prefer losers to winners. Anim. Behav., 66: 399-407.

SCHLEIDT, W.M. AND SHALTER, M.D., 1973. Stereotypy of a fixed action pattern during ontogeny in Coturnix coturnix coturnix. Z. Tierpsychol., 33: 35-37.

SEZER, M., BERBEROGLU, E. AND ULUTAS, Z., 2006. Genetic association between sexual maturity and weekly live- weights in laying-type Japanese quail. S. Afr. J. Anim. Sci., 36: 142-148.

SHARP, S.P. AND HATCHWELL, B.J., 2005. Individuality in the contact calls of cooperatively breeding long-tailed tits (Aegithalos caudatus). Behaviour, 142: 1559-1575.

(Received 10 March 2010, revised 29 March 2010)

M. Sezer* and O. Tekelioglu Karamanoglu Mehmetbey Universitesi, Meslek Yuksekokulu, Ev ve Deney Hayvanlari Programi, Karaman,Turkey (MS), and Tunceli Universitesi, Muhendislik Fakultesi, Makina Muhendisligi Bolumu, Tunceli, Turkey (OT) Corresponding author: msezer@kmu.edu.tr 0030-9923/2010/0005-0581 $ 8.00/0 Copyright 2010 Zoological Society of Pakistan.
COPYRIGHT 2010 Asianet-Pakistan
No portion of this article can be reproduced without the express written permission from the copyright holder.
Copyright 2010 Gale, Cengage Learning. All rights reserved.

Article Details
Printer friendly Cite/link Email Feedback
Author:Sezer, M.; Tekelioglu, O.
Publication:Pakistan Journal of Zoology
Article Type:Report
Geographic Code:7TURK
Date:Oct 31, 2010
Words:3139
Previous Article:Forecasting of Spotted Bollworm (Earias vitella (Fab.) (Lepidoptera: Noctuidae) Occurrence in Cotton.
Next Article:Characterization of Multiple Antimicrobial Resistant Shigella sonnei Isolated from Diarrhoeal Patients in Azad Kashmir, Pakistan*.
Topics:

Terms of use | Privacy policy | Copyright © 2021 Farlex, Inc. | Feedback | For webmasters |