Meta-analysis: Supplementary artificial light and goose reproduction

Journal Pre-proof Meta-analysis: Supplementary artificial light and goose reproduction G.J. Liu, Z.F. Chen, X.H. Zhao, M.Y. Li, Z.H. Guo

PII:

S0378-4320(19)30849-8

DOI:

https://doi.org/10.1016/j.anireprosci.2020.106278

Reference:

ANIREP 106278

To appear in:

Animal Reproduction Science

Received Date:

9 September 2019

Revised Date:

20 December 2019

Accepted Date:

7 January 2020

Please cite this article as: Liu GJ, Chen ZF, Zhao XH, Li MY, Guo ZH, Meta-analysis: Supplementary artificial light and goose reproduction, Animal Reproduction Science (2020), doi: https://doi.org/10.1016/j.anireprosci.2020.106278

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Meta-analysis: Supplementary artificial light and goose reproduction G. J. Liua *, Z. F. Chenb, #, X. H. Zhao a, M. Y. Li a, Z. H. Guoa,*

a

Heilongjiang Academy of Agricultural Sciences, Animal Husbandry Research

Institute, No. 368 Xuefu Road, Harbin 150086, P. R. China b

Heilongjiang Animal Sciences Institute, No. 2 Heyi Road, Qiqihare 161005, P. R.

China *

Co-corresponding author: G. J. Liu, Heilongjiang Academy of Agricultural Sciences

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(HAAS), Animal Husbandry Research Institute, No. 368 Xuefu Road, Harbin 150086, P. R. China. E-mail: [email protected], FAX/ TEL: 86-451-87502330 *

Co-corresponding author: Z. H. Guo, Heilongjiang Academy of Agricultural Sciences

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(HAAS), Animal Husbandry Research Institute, No. 368 Xuefu Road, Harbin 150086,

#

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P. R. China. E-mail: [email protected], FAX/ TEL: 86-451-87502330

G. J. Liu and Z. F. Chen contributed equally to this work and are considered equal first

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authors Z. F. Chen, E-mail: [email protected]

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X. H. Zhao , E-mail: [email protected]

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M. Y. Li , E-mail: [email protected]

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ABSTRACT

Photoperiod affects poultry reproduction, and in birds, photoperiod regulation is a

complex physiological process. In modern poultry production, lighting management has become an important and effective management approach for increasing egg production. Geese are domesticated fowl and in many goose production enterprises animals are allowed to roam in outside pens during the day and are housed indoors at night, so the animals can be exposed to artificial lighting during the night periods.

Supplementary artificial lighting resulted in improved reproduction in some studies, but reports have been inconsistent. To evaluate the results from previous studies of supplementary lighting on goose egg production, a meta-analysis was conducted to determine optimal supplementary artificial lighting regimens for geese egg production. Results indicated supplementary artificial light increases mean egg production, the length of the period of egg production before there is cessation of egg production capacity, and fertility. In summary, there were evaluations of data from five studies focused on White Roman geese in the meta-analysis conducted in the present study, however, examination of more breeds is necessary to make more definitive assessments

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of the findings from this meta-analysis.

Keywords: Egg number; Geese; Laying; Light

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1. Introduction

There is a long tradition of goose breeding in in Europe and China (Ren et al.,

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2016). Photoperiod adjustments are used during the reproduction period to enhance egg production (Zeman et al., 1990; Wang et al., 2009; Chang et al., 2016a). In different

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regions of the world, there is use of different lighting regimens to modulate the reproductive performance when taking into consideration latitude where the birds are located and the different geese breeds that are predominant in the different regions. In

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the south of mainland China, during the period preceding onset of egg production, there is imposing of an 18 hour light:6 hour dark photoperiodic regimen (18L:6D) for a 2 to

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2.5 month period, followed by an 11L:13D photoperiodic regimen during the egg production period (Chang et al., 2016a). In Taiwan, geese are reared using a regimen of 7L:17D photoperiodic regimen for 6 weeks, followed by a 9L:15D photoperiodic

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regimen during the egg-production period (Chang et al., 2016c). The lighting programme in Europe starts about 30 days before the season of egg production by imposing of a 9L:15D photoperiodic regimen, then the light period is extended to 10 to 12 hours per 24-hour period during the period of egg production (Rosinski, 2019). In different regions of the world, there are different goose breeding seasons preferred, so many of these practices are latitude-dependent (Wang et al., 2002a). Although the origins of the domestic geese are unclear (Heikkinen et al., 2015), results of one study

indicate there were two regions in which domestication of Chinese and European geese occurred (Wang et al., 2017). It is likely that controlling lighting regimens can be utilized for improving goose reproduction, specifically egg production (Chang et al., 2016d). Although artificial lighting can easily be used to manipulate the timing of egglaying periods in windowless buildings, the large economical costs of this type of facility render it impractical in many situations (Wang et al., 2005). With the invention of the light emitting diode technology (LED), it has become economically feasible to build goose housing facilities and include the LED technology in the physical infrastructure of buildings (Chang et al., 2016c).

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Results of one study indicate the use of supplementary artificial light leads to an improved goose reproduction, specifically egg production (Wang et al., 2009), but other results indicate there are not beneficial effects of artificial lighting in goose production enterprises (Wang et al., 2002b; Wang et al., 2005). For this reason, we conducted a meta-analysis to ascertain whether artificial light can affect the efficiency of goose

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reproduction.

2.1. Database and data extraction

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2. Methods

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Electronic searches were conducted to identify relevant articles and papers. These were selected by two authors independently using PubMed, Ovid, ProQuest, and

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ScienceDirect from January 1, 2000 to July 1, 2019 using the key words: (goose OR geese OR goosie OR gooses OR goslings OR anser) AND (laying OR egg-laying OR (egg number) OR (egg No.)) AND (light OR lighting OR photoperiod). Limiting

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categorical terms were used in titles, abstracts, and keywords. Information in Table 1 provides the study inclusion and exclusion criteria used in this study.

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2.2. Meta-analysis

There was collection of data regarding supplementary artificial light for goose egg

number, duration of egg production period before cessation of production occurs, and fertility. When data required calculation, there was a modified standard error of the mean (SEM) to a standard deviation (SD). There was determination of whether there is heterogeneity of results using the Higgins statistic and the P-value and I2 statistic (I2 >50% indicated a large amount of heterogeneity) (de la Cruz et al., 2017). All heterogeneity analyses that were conducted, and subgroup analysis procedures that

were conducted are reported in Table 2. Data analyses were conducted using Review Manager (Version 5.3, Copenhagen: Nordic Cochrane Centre, Cochrane Collaboration) utilizing a fixed-effect model because all geese were White Roman. A continuous model was used to assess data for supplementary artificial light, number of eggs, and duration of the egg production period before there was cessation of production. A dichotomous model was used to assess fertility. Publication bias was assessed using Begg funnel plots, Egger’s test (Egger et al., 1997), and Begg’s test (Stata Corp 12.0, College Station, TX, US).

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3. Results Records (n = 97) were identified in the initial electronic search. After assessing the information provided in each study, five studies were included in the meta-analysis

(Fig. 1) (Wang et al., 2002a; Wang et al., 2002b; Wang et al., 2005; Wang et al., 2009; Chang et al., 2016a). The main characteristics of the studies are provided in Table 2.

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Results were pooled, and Figures 2, 3, and 4 depict that with use of supplementary artificial light there is an increase in the number of eggs produced (95% CI, 0.44–7.09),

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duration of the egg production period before cessation of egg laying occurs (95% CI, 2.49–22.93), and fertility (95% CI, 1.16–1.43). Heterogeneity was confirmed, as

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indicated by information included in Table 2. The factors affecting the response to supplementary artificial lighting are the intensity of illumination, sex ratios, and age. When there was subgroup analysis for illumination intensity at 50 and 100 lux (Fig. 2),

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when there were ganders housed with the female geese (Fig. 3) and birds were older than 2 years of age (Fig. 4), there was a lack of data heterogeneity. Consequently, there

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was recalculation of the means from the studies (Table 3). With the provision of supplementary artificial lighting, this resulted in an increase in mean egg numbers, period of duration of egg production before there was cessation of production, and

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fertility. Risk of bias is depicted in Figure 5, and there was no publication bias detected with the meta-analysis of the data in this study. When the Egger’s (P = 0.689) and Begg’s tests (Pr > |z| = 0.806) were also used to evaluate bias, there was confirmation that there was not bias of the data evaluated using the meta-analysis. 4. Discussion 4.1. Main finding

The use of supplementary artificial lighting in goose egg production results in an increase in mean egg number produced by each female, duration of the egg layingperiod without cessation of egg production, and fertility rate when ganders are housed with the female geese for purposes of production of fertilized eggs. Light directly and indirectly through dietary intake provides energy for living organisms and affects their growth, development, and reproduction (Penteriani and Delgado, 2017). Photoreceptors that induce the response to different avian photoperiods are located in the head rather than in the eye as occurs in most other animals (Underwood and Menaker, 1970). Light activates photoreceptors in the hypothalamus,

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where the light signals are transformed into neural signals and ultimately endocrine signals that are transferred to various body tissues (Follett et al., 1977). If the period

during which supplementary artificial lighting is provided lasts too long,

photorefractoriness occurs (Siopes, 1997). Type I, II and IV cryptochromes (Cry1, Cry2,

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and Cry4) are present in the eyes of birds. The Cry1 and Cry2 proteins may be encoded

by circadian clock genes (Watari et al., 2012), but the patterns of Cry4 gene expression are not consistent with the light patterns resulting from photoperiodic regimens; rather

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it has been speculated that the production of these proteins may be part of the magnetic compass of birds (Pinzon-Rodriguez et al., 2018). Results of one study indicated that

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birds perceive signals through their categorical perception of different colours (Caves et al., 2018).

Deep-brain photoreceptors (DBP) are located in the paraventricular organ (PVO)

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and contain opsins and neuropsins that can receive lighting stimuli that modulates the functions of these receptors (Nakane et al., 2010). There is a neural response to the

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lighting stimuli that is transmitted via specialized neurons to the pars tuberalis (PT), where there is release of thyroid stimulating hormone (TSH) induced. The TSH subsequently induces the activation of the type 2 deiodinase (Dio2). The Dio2 functions

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to stimulate the release of gonadotropin-releasing hormone (GnRH) (Zhu et al., 2019a). The GnRH subsequently stimulates the release of luteinizing hormone (LH) and follicle-stimulating hormone (FSH) from the anterior pituitary (Zhu et al., 2019b). Melatonin secretion of the blackheaded bunting (Emberiza melanocephala) was decreased when there was light stimulation (Malik et al., 2015). Melatonin is mainly synthesized by pineal cells with synthesis and secretion being regulated by photoperiodic signalling with there being circadian and seasonal changes in pattern of

secretion. The melatonin receptor belongs to the G protein coupled receptor superfamily. There are three subtypes of MT1, MT2 and MT3 in mammals. Three subtypes of Mel 1a, Mel 1b and mel-1c were found to be present in poultry (Dubocovich et al., 2010). Seasonal goose breeding and large viable broods are primary factors affecting the reproductive performance and period of egg production in geese (Barnas et al., 2018). The goose breeding season lasts from October to March in Israel (Pyrzak et al., 1984), from October to May in Taiwan (Wang et al., 2009) and from February to June in Europe (Rosinski, 2019). Goose eggs are larger than hen eggs, usually weighing 125 to 160 g (Wang et al., 2002a), thus, there is a greater physiological energetic requirement

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for producing eggs and this factor affects the number of eggs produced. The use of supplementary artificial lighting results in a delay when the first egg is produced as compared to when there is no supplementary lighting imposed (Wang et al., 2002a).

Goose reproductive capacity is regulated by the functions of the hypothalamic–

pituitary–gonadal axis (Liu et al., 2018). Candidate functional genes related to goose

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egg-laying include follicle stimulating hormone (Du et al., 2018), gonadotropinreleasing hormone (Hirschenhauser et al., 2000), pituitary-specific transcription factor-

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1 (Liu et al., 2008), and prolactin (Chen et al., 2012). The photoperiod affects gonadal and serotonergic axes in ganders by functioning to modulate gene expression regulation

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and hormone secretion (Gumulka and Rozenboim, 2015). The photoperiod affects expression of specific genes in geese. Results from a recent study indicate goose vasoactive intestinal peptide (VIP) mRNA transcript

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abundance was less when there was imposing of a short, 8L:16D, as compared to the 11L:13D photoperiodic treatment regimen. The results from the same study indicated prolactin (PRL) mRNA transcript abundance was also lesser when there was imposing

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of the 8L:16D photoperiodic regimen as compared with the 11L:13D photoperiodic treatment regimen. The results of the study indicate that the imposing of the 11L:13D

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photoperiodic regimen resulted in a longer period of egg production before cessation of egg laying as compared with the other photoperiodic regimens assessed (Zhu et al., 2019b).

The colour of the light also affects the egg-laying rate in geese. Results of one recent study indicate that the use of white and red LED lighting when imposing an 11L:13D photoperiodic regimen can increase the egg-production rate of geese. There was greater geese egg production with these colour lighting regimens than with blue and green light treatments, because of differences in PRL, VIP, and gonadotropin-

inhibitory hormone (GnIH) gene expression with these lighting regimens (Zhu et al., 2019a). Light intensity is the most important photoperiodic factor affecting bird physiology and there are differential effects when there are differing light intensities that are dependent on light perception by the hypothalamic photoreceptors (Rozenboim et al., 2013). Supplemental light intensity has a marked effect on goose egg production (Pyrzak et al., 1984). With use of supplementary lighting that has a 170 lux light intensity, egg-laying and egg weights of White Roman geese were greater than that of the control birds (Chang et al., 2016b). Thus, the intensity of the light is a source of

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heterogeneity in studies of supplementary artificial light and number of eggs produced. The reproductive performance of hens (turkey, ducks, and quail) depends on the number of eggs produced and whether the female reproductive behavior is conducive

to mating (Ottinger et al., 1982; Raud and Faure, 1990). Males often selectively mate

with specific females, and females can also control sexual activity via complex

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physiological and behavioral characteristics (Cezilly et al., 2000). Whether the ratio of

males to females affects numbers of eggs produced, however, is not clear. With

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evaluation of the results from the meta-analysis of the present study, sex ratios in the groups of egg-producing female geese were an apparent source of heterogeneity when

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the duration egg production was evaluated. Goose age also effects mating behavior and fertilization (Gumulka and Rozenboim, 2013). Age is a source of heterogeneity in egg production even when there is supplementary artificial lighting regimens used to

4.2. Limitations

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increase fertility.

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The laying period for White Roman geese is in June and July (Wang et al., 2002b). The laying period of local hybrid geese in Israel is from October to June of year

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following when there was hatching of these females (Pyrzak et al., 1984). Seasonal goose reproduction may be related to latitude (Wang et al., 2005). The five studies evaluated with the meta-analysis in the present study focused on White Roman geese, and results indicated that prolonged artificial lighting may enhance reproduction, but studies in other breeds are required to further assess these findings. 4.2. Implications

Supplementary artificial light increases goose reproduction, but more research is needed to more precisely determine specific effects of lighting regimens and for development of more effective lighting regimens for enhancing reproduction. Acknowledgments We thank LetPub (www.letpub.com) for its linguistic assistance during the preparation of this manuscript.

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Author Statement G. J. Liu and Z. H. Guo design the idea.

X. H. Zhao and M. Y. Li made electronic searches at internet,

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When the study inclusion and exclusion were difficult to decide, Z. F. Chen make final decision.

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Z. H. Guo and Z. F. Chen procedure meta-analysis.

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G. J. Liu and Z. H. Guo work out the draft.

Funding

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This research was supported by the Harbin Science and Technology Innovation Talents Project (Grant number 2017 RAXYJ021); National Waterfowl Industry

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Technology System Project (Grant number CARS-42-24). Conflicts of interest

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None of the authors have any conflicts of interest.

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Tables Table 1 Inclusion and exclusion criteria Exclusion No geese evaluated Non-English No light No egg number data Review

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Inclusion Animals and poultry including but not limited to geese English Light but not limited to light Egg number data included Original research

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Characteristics of studies included in the review

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Table 2

Studies

Light:Dark

Illumination intensity (lux)

Crude protein

1 2 3 4 5

Wang 2002b Wang 2002a Wang 2005 Wang 2009 Chang 2016

20L:4D 16L:8D 19L:5D 13.5L:10.5D 19L:5D

40–50 40–50 120 53 30–40

18% 18% 18% 18% 15% or 18%

e-

Pr

na l Jo ur

pr

No.

Metabolizable energy (MJ ME/kg) 11.51 11.51 11.21 11.21 11.09

Age

Sex ratio ganders: geese

9.5 months 8 months 18 months 1 year 2.7 years

No ganders 6:16 1:4 1:4 2:5

f oo

Table 3

Control

32.83±6.99 10.34

Pr

Supplementary artificial light

Jo ur

na l

Rise percent (%)

Laying duration

e-

egg number 29.44±4.72

pr

Average data from five studies

egg fertility rate

92.3±19.3

40.03±6.52

118.02±26.91 21.79

47.44±7.26 15.63

Figure Legends

na

lP

re

-p

ro of

Fig. 1. Flow diagram of the meta-analysis depicting each study

Fig. 2. Forest plot of the effect of supplementary artificial light effect on number of

Jo

ur

eggs produced; CI = 95% confidence interval

ro of

Fig. 3. Forest plot of the effect of supplementary artificial light on duration of egg

na

lP

re

-p

production period; CI = 95% confidence interval

ur

Fig. 4. Forest plot of the effect of supplementary artificial light on fertility; CI = 95%

Jo

confidence interval

ro of

Fig. 5. Funnel plots of the effect of supplementary artificial light on number of eggs

produced; Empty circles indicate less than 50 lux, Empty squares indicate more than 50

Jo

ur

na

lP

re

-p

lux