Effect of thermal conditioning on growth performance and thermotolerance in broilers: A systematic review and meta-analysis.

✅ 全文

热调节对肉鸡生长性能和耐热力的影响:系统综述与荟萃分析

作者 C. Ncho; Vaishali Gupta; A. Goel 期刊 Journal of thermal biology 发表日期 2021 DOI 10.1016/j.jtherbio.2021.102916 类型 原创研究 (Original Research)

📄 中文摘要 Chinese Abstract

中文
热 conditioning 已被引入作为提高肉鸡生长性能和耐热性的成本效益策略。然而,由于各研究之间实验条件的差异性,得出一致的结论具有挑战性。本系统综述和荟萃分析旨在量化早期热 conditioning 对肉鸡关键生产和生理参数的影响。分析重点为热中性条件下的生长性能以及热应激下的体温反应。

📋 英文结构化总结 English Structured Summary

全文整理

EN

Background:

Thermal conditioning has been introduced as a cost-effective strategy to enhance growth performance and thermotolerance in broilers. However, due to variability in experimental conditions across studies, drawing consistent conclusions is challenging. This systematic review and meta-analysis aimed to quantify the effects of early-age thermal conditioning on key production and physiological parameters in broiler chickens. The analysis focused on growth performance under thermoneutral conditions and body temperature responses under heat stress.

Methods:

A comprehensive literature search was conducted in December 2020 across Scopus, PubMed, Scielo, Web of Science, and Google Scholar using keywords related to thermal conditioning and broilers. Studies were included if they involved post-hatch heat conditioning in broilers, reported relevant outcomes (body weight gain, feed intake, feed conversion ratio, body temperature), and provided sufficient statistical data. A restricted maximum likelihood random-effects model was used to pool effect sizes, with standardized mean differences (SMD) for most traits and mean differences (MD) for feed conversion ratio. Subgroup analyses were performed based on heat stress type (acute vs. chronic), and heterogeneity and publication bias were assessed.

Results:

Thermal conditioning significantly increased body weight gain (SMD = 0.139, 95% CI: 0.0372–0.2407, *P* = 0.0074) and feed intake (SMD = 0.292, 95% CI: 0.108–0.476, *P* = 0.0019) under thermoneutral conditions compared to controls. Feed conversion ratio was not significantly affected (MD = −0.013, 95% CI: −0.0307 to 0.0041, *P* = 0.1346). Body temperature was significantly reduced under acute heat stress (SMD = −0.455, 95% CI: −0.718 to −0.192, *P* < 0.001), but no significant effect was observed under chronic heat stress (SMD = −0.115, 95% CI: −0.651 to 0.420, *P* = 0.6729).

Data Summary:

The meta-analysis included 17 studies. For body weight gain, data from 50 groups showed a mean of 1702.2 g (range: 94.7–2849.5 g); feed intake from 48 groups averaged 30,005.8 g (range: 37.7–4940 g); feed conversion ratio from 40 groups had a mean of 1.92 (range: 1.38–2.7); and body temperature from 57 groups averaged 43.2°C (range: 40.32–46.37°C). Heterogeneity was moderate to high across all outcomes (I² > 57%).

Conclusions:

Early-age thermal conditioning improves body weight gain and feed intake in broilers under thermoneutral conditions and effectively reduces body temperature during acute heat stress. However, it does not significantly affect feed conversion ratio or confer thermotolerance under chronic heat stress. These findings support the use of thermal conditioning as a practical tool to enhance productivity and acute heat resilience in broiler production.

Practical Significance:

Thermal conditioning can be implemented in commercial broiler farming as a low-cost, non-nutritional intervention to improve growth performance and prepare birds for acute heat events, thereby supporting sustainable poultry production in warming climates.

📋 中文结构化总结 Chinese Structured Summary

中文

背景:

热 conditioning 已被引入作为提高肉鸡生长性能和耐热性的成本效益策略。然而,由于各研究之间实验条件的差异性,得出一致的结论具有挑战性。本系统综述和荟萃分析旨在量化早期热 conditioning 对肉鸡关键生产和生理参数的影响。分析重点为热中性条件下的生长性能以及热应激下的体温反应。

方法:

于2020年12月在Scopus、PubMed、Scielo、Web of Science和Google Scholar中进行了全面的文献检索,使用与热 conditioning 和肉鸡相关的关键词。纳入标准包括:涉及肉鸡孵化后热 conditioning、报告相关结果(体重增加、采食量、饲料转化率、体温)并提供充分的统计学数据。采用限制性最大似然随机效应模型合并效应量,大多数性状使用标准化均差(SMD),饲料转化率使用均差(MD)。根据热应激类型(急性与慢性)进行亚组分析,并评估异质性和发表偏倚。

结果:

与对照组相比,热 conditioning 在热中性条件下显著提高了体重增加(SMD = 0.139,95% CI:0.0372–0.2407,*P* = 0.0074)和采食量(SMD = 0.292,95% CI:0.108–0.476,*P* = 0.0019)。饲料转化率未受到显著影响(MD = −0.013,95% CI:−0.0307至0.0041,*P* = 0.1346)。在急性热应激下体温显著降低(SMD = −0.455,95% CI:−0.718至−0.192,*P* < 0.001),但在慢性热应激下未观察到显著效应(SMD = −0.115,95% CI:−0.651至0.420,*P* = 0.6729)。

数据汇总:

本荟萃分析共纳入17项研究。体重增加方面,50组数据的均值为1702.2 g(范围:94.7–2849.5 g);采食量方面,48组数据的均值为30,005.8 g(范围:37.7–4940 g);饲料转化率方面,40组数据的均值为1.92(范围:1.38–2.7);体温方面,57组数据的均值为43.2°C(范围:40.32–46.37°C)。所有结局指标的异质性为中等至高度(I² > 57%)。

结论:

早期热 conditioning 可改善热中性条件下肉鸡的体重增加和采食量,并有效降低急性热应激期间的体温。然而,它对饲料转化率无显著影响,也未在慢性热应激下赋予耐热性。这些发现支持将热 conditioning 作为提高肉鸡生产力和急性热应激恢复力的实用工具。

实践意义:

热 conditioning 可在商品肉鸡养殖中作为一种低成本、非营养性干预措施实施,以改善生长性能并使鸡只做好应对急性热事件的准备,从而在气候变暖背景下支持可持续的家禽生产。

📖 英文全文 English Full Text

EN

Journal of Thermal Biology 98 (2021) 102916 Available online 25 March 2021

0306-4565/© 2021 Elsevier Ltd. All rights reserved.

Effect of thermal conditioning on growth performance and thermotolerance in broilers: A systematic review and meta-analysis

Chris Major Ncho a, Vaishali Gupta b, Akshat Goel a,* a Department of Animal Science, Gyeongsang National University, Jinju, 52828, Republic of Korea b Department of Applied Life Science, Gyeongsang National University, Jinju, 52828, Republic of Korea

A R T I C L E I N F O Keywords:

Thermal conditioning Broilers Heat stress Meta-analysis

A B S T R A C T Thermal conditioning has been introduced as a cost-effective way to improve performance and thermotolerance in broilers. However, since all the trials were performed under various experimental conditions, it appears difficult to draw general conclusions. Therefore, the objective of this study was to quantify the response of broilers to thermal conditioning through a meta-analysis approach. A literature search was conducted on Scopus,

PubMed, Scielo, Web of Science, and Google scholar in December 2020. A restricted maximum likelihood random effect model was used to pool the effect sizes from the body weight gain (BWG), feed intake (FI), feed conversion ratio (FCR), and body temperature (Tb). BWG, FI, and Tb were computed using the standardized mean difference (SMD) while FCR was computed using mean differences (MD) with a 95% confidence interval (IC). Growth performances were evaluated during the thermoneutral conditions while Tb was evaluated after either acute or chronic heat stress after early age thermal conditioning. A total of 17 studies were included in the dataset. Thermal conditioning significantly increased BWG (SMD = 0.139, IC = 0.0372–0.2407, P = 0.0074) and

FI (SMD = 0.292, IC = 0.108–0.476, P = 0.0019) compared with the control. Additionally, subgroup analysis revealed that overall Tb was significantly reduced under acute heat stress (SMD = -0.455, IC = -0.718 to −0.192,

P < 0.001) but not affected during chronic heat stress (SMD = -0.115, IC = -0.651 to −0.420, P = 0.6729). In conclusion, thermal conditioning significantly increased the broiler’s BWG and FI under thermoneutral condi­ tions and can help in reducing Tb under acute heat stress.

1. Introduction As the global climate changes, environmental variation becomes a challenge for animal production (Nawab et al., 2018). Particularly in poultry cultivation, modern broiler breeds have been extensively sub­ mitted to selection for increasing their growth performances under mild climates (Sandercock et al., 2006). Additionally, the absence of sweat glands, as well as the presence of feathers, implies that chickens cannot regulate body temperature in high-temperature environments (Xie et al.,

2015).

Several studies demonstrated that early age thermal conditioning can be considered as a technique for mitigating the adverse impact of high ambient temperature in broiler production (De Basilio et al., 2001;

Oke et al., 2020; Yahav, 1999). Although different temperatures and time-period have been tried by the researchers, the method for thermal conditioning is similar and the temperature is kept slightly higher than the recommended one for few hours or even days for this purpose (Fig. 1). The improvement of broiler’s thermotolerance via thermal conditioning was made possible due to the immaturity of the mechanism of temperature regulation during the first week of their life (Modrey and

Nichelmann, 1992) which is firmly linked to sympathetic neural activity and the integration of thermal information in the hypothalamus (Kin­ ney, 1992). In addition, evidence shows that submitting broilers to thermal conditioning can bring about better development in growth performances as well as higher carcass yield (Bengharbi et al., 2016b;

Uni et al., 2001).

However, despite the fact that extensive research has been conducted to explore the role of thermal conditioning, there are numerous di­ vergences among the outcomes acquired from the literature. Therefore, it is critical to evaluate the previously published data to discover the potential effectiveness of thermal conditioning in broiler production.

The current study aimed to quantify the reaction of broilers to thermal conditioning through a meta-analysis approach. Growth parameters such as body weight, feed intake, and feed conversion ratio were

* Corresponding author.

E-mail address: genesakshat@gmail.com (A. Goel).

Contents lists available at ScienceDirect Journal of Thermal Biology journal homepage: http://www.elsevier.com/locate/jtherbio https://doi.org/10.1016/j.jtherbio.2021.102916

Received 3 February 2021; Received in revised form 19 March 2021; Accepted 19 March 2021

Journal of Thermal Biology 98 (2021) 102916 2 evaluated during thermoneutral conditions and body temperature was evaluated during high ambient temperature conditions.

2. Materials and methods 2.1. Search and data filtering

The studies’ research was conducted by consulting the following electronic databases: Scopus, PubMed, Scielo, Web of Science, and

Google scholar in December 2020. Performing the research in such a wide variety of repositories prevents from including articles found in only one database. The keywords considered for the research were:

“thermal acclimation”, “thermal conditioning”, “heat conditioning”,

“temperature conditioning”, “broilers” as well as their different combi­ nations. Two independent researchers conducted the research. Studies before inclusion in the database were critically analyzed subsequently by their title, abstract, and finally the full text. A study was selected only after the agreement of both researchers. No author has been contacted to ascertain further information or to obtain unpublished data.

The main criteria to select the published articles were: (i) articles should be published in the English language in peer-reviewed journals; (ii) broilers were used as experimental animals; (iii) articles should include at least one treatment of thermal post-hatch conditioning; (iv) all thermal conditioning parameters (temperature, duration, starting age) should be available in the material and method; (v) articles should evaluate heat conditioning (cold conditioning studies were not included in the database); (vi) for studies evaluating heat stress responses there should be a control treatment under heat stress conditions. After the filtering step, the final database included data collected from 17 papers.

2.2. Data recording and coding The response traits extracted from the studies were categorized as follows: growth parameters and body temperature. From each study, the mean, standard deviation, or standard error of the mean (SEM) and the number of animals in each treatment were extracted. When the standard deviation was not reported, it was calculated based on the standard error of the mean. Since there was a shortage of data from studies that reported growth performances under heat stress and to just consider the effect of the thermal conditioning, only the growth parameters reported under standard housing temperature were included in the dataset. The body weight gain (BWG), feed intake (FI), and feed conversion ratio (FCR) were the traits considered. The body temperature under heat challenge (Tb) was the last response trait extracted for analysis.

2.3. Statistical analysis The package “metafor” of the R software version 4.0.3 (R Core Team,

2020), was utilized to conduct all statistical analyses and generate fig­ ures. A restricted maximum likelihood random effect model was used to analyze the dataset. Due to the distinct nature of the response traits, they were computed using the standardized mean difference (SMD) except

FCR which was transformed in the mean difference (MD) as previously executed by De Toledo et al. (2020). Concerning the Tb, a subgroup analysis was carried out by splitting the different heat stress categories to evaluate heterogeneous results and making comparisons between these groups. The two subgroups considered were chronic and acute heat stress. Studies were included in one group or another depending on the category of heat stress they considered in their design. Heterogeneity was evaluated by the calculation of the Chi-square test (Q) and the I2 statistic. Substantial heterogeneity was noticed if P-value < 0.05 (Q) and/or I2 > 50%. Finally, publication bias was assessed via Egger’s test and the realization of funnel plots.

3. Result 3.1. Study selection process Details regarding the selection process of studies are showed in

Fig. 2. The literature searches in the electronic repositories for articles published from 1996 to 2020 retrieved a total of 8040 studies. After removing 1348 duplicates, the number of articles remained 6692. A thorough screening of the abstracts and the titles of these articles resulted in the exclusion of 6662 records that were not directly linked to our topic. Subsequently, from the 30 articles retained, 11 were excluded due to the lack of significant data helpful in the study after analysis of the full text. Moreover, 2 articles that were evaluating cold conditioning were not selected in light of the fact that the motivation behind this review was just to access the impacts of heat conditioning.

3.2. Dataset description Table 1 shows the characteristic of the articles selected for the meta- analysis. The frequency of strains used in the studies was the following:

Arbor Acres (n = 1); Ross (n = 3); Hubbard (n = 3); Cobb (n = 7) and 3 studies did not specify the strain of the broiler chicken. Fig. 3 presents the bubble chart of the studies selected in the meta-analysis. The tem­ perature of conditioning varied widely from 35 ◦C to 40.5 ◦C. Most of the studies applied a duration of conditioning of 1 day (n = 11). A total of 4 studies did not include a heat challenge in their design while 6 opted for chronic heat stress and 7 for acute heat stress. The overall heat stress temperature variation ranged from 30 ◦C to 43 ◦C. The most-reported response trait was BWG (14 studies) followed by FI (12 of studies),

FCR (9 studies), and Tb (9 studies). The summary statistics of the re­ sponses traits evaluated are presented in Table 2.

3.3. The effect of thermal conditioning on growth performances in broilers

The forest plot (Fig. 4) indicates that there was an overall positive effect on the BWG of birds that were thermally conditioned compared to the control birds (SMD = 0.139, IC = 0.0372–0.2407, P = 0.0074).

Heterogeneity was present in this effect since I2 = 57.7% with P-value

<0.001. Similarly, in Fig. 5 the diamond position reflects the positive effect of thermal conditioning on the FI in broilers. The overall effect

Fig. 1. Diagram explaining thermal conditioning process. Thermal condition­ ing consists of exposing birds in their first week of life to acute heat spells in order to enhance their thermal resistance in their later life. Temperature can range from 35 ◦C up to 40.5 ◦C and the duration of exposure is usually 24 h.

C.M. Ncho et al.

Journal of Thermal Biology 98 (2021) 102916 3 was estimated at SMD = 0.292, IC = 0.108–0.476, P = 0.0019 with significant heterogeneity (I2 = 85.7% with P-value <0.001). Fig. 6 shows that there was no significant effect of thermal conditioning on the

FCR of broilers (MD = -0.013, IC = -0.0307–0.0041, P = 0.1346).

Moreover, significant heterogeneity was observed (I2 = 89.7%; P value

< 0.001).

3.4. The effect of thermal conditioning on body temperature in broilers

During the heat challenges, the difference in body temperature be­ tween the thermally conditioned birds and the control was wider at the two extreme ambient temperatures (30 and 43 ◦C) while between 36 and

39 ◦C, body temperatures of both groups were similar (Fig. 7A). During acute heat stress, the birds recorded an overall higher body temperature compared to the data collected from studies that evaluated chronic heat stress (Fig. 7B). The diamond in the forest plot (Fig. 8) indicates an overall lower Tb in thermally conditioned birds compared to the control group (SMD = -0.315, IC = -0.584 to −0.046, P = 0.0219) under heat stress. This effect was significantly heterogeneous (I2 = 73.9% with P- value <0.001). The subgroup analysis showed a significant reduction in the body temperature of thermally conditioned birds during acute heat stress (SMD = -0.455, IC = -0.718 to −0.192, P < 0.001). However, under chronic heat stress, thermal conditioning did not show any statistically significant difference in Tb compared to the control group (SMD = -0.115, IC = -0.651 to −0.420, P = 0.6729). Publication bias.

Fig. 9 shows funnel plots of the parameters studied in this meta- analysis. The symmetry of all funnel plots was confirmed by Egger’s linear regression test. Results were not significant for BWG (t = 0.06, df

= 31, P = 0.9524), FI (t = -0.40, df = 29, P = 0.6925), FCR (t = 0.65, df

= 24, P = 0.5245), and Tb (t = 0.84, df = 30, P = 0.4098). Thus, there was no risk of publication bias detected in this study.

4. Discussion The livestock industry is extremely competitive. Extensive selection and nutritional programming emphasize for identifications of new methods for improving growth performances and heat tolerance in broiler chickens. Several trials evaluating the impact of early thermal conditioning on growth performances and thermotolerance of broilers have been conducted over the last decade. Nonetheless, up-to-date in­ formation summarizing the result from the previous experiment on this topic remains scarce. Thus, this review was an attempt to evaluate the potential outcomes of thermal conditioning in broilers through a sta­ tistical meta-analysis approach. In this study, since moderate to high heterogeneity was detected in all the response traits evaluated, a random effect model was applied. Indeed, random-effect methods

Fig. 2. Process for studies selection. A total of 8040 studies were retrieved from the electronic repositories (PubMed, Web of Science, Scopus, Scielo, Google Scholar).

Finally, a total of 17 studies were included in the meta-analysis.

C.M. Ncho et al.

Journal of Thermal Biology 98 (2021) 102916 4 explicitly account for the heterogeneity which represents the across- study variation (DerSimonian and Kacker, 2007).

One of the primary after-effects of the current examination is that thermal conditioning significantly increased the broiler’s BWG and FI while FCR was not affected. In general, the short-term exposure to heat during thermal conditioning results in subsequent growth retardation (Yahav and Hurwitz, 1996; Yahav et al., 1997). It is believed that the final increase in BWG and FI of thermally conditioned broilers is the result of a compensatory growth similar to that observed during feed restriction at an early age (Plavnik and Hurwitz, 1985). Indeed, it has been acknowledged that high housing temperature causes a deprivation in broiler’s FI leading to a reduction in body weight (Awad et al., 2020;

Farag and Alagawany, 2018). Therefore, growth is accelerated post-conditioning to totally compensate for the losses and sometimes result in higher body weight at marketable age (Yahav and McMurtry,

Table 1 Details of the studies included in the meta-analysis.

Author and year Strain Response variable extracted

Heat challenge Oke et al. (2020) Arbor acres BWG,FCR,FI

None Zaboli et al. (2017) Ross 308 BWG,FCR,FI,Tb Chronic

Uni et al. (2001) Cobb BWG,FCR,FI None De Basilio et al. (2001)

Cobb BWG,FI,Tb Acute Yahav and McMurtry (2001) Cobb

BWG,FCR,Tb Acute Taouis et al. (2002) Non- specified

BWG None Liew et al. (2003) Cobb BWG Chronic Yahav et al. (1997)

Non- specified BWG,FI Acute Abdelqader and Al-Fataftah (2014)

Hubbard BWG,FI,Tb Acute Kang et al. (2019) Ross BWG,FCR,FI

Chronic Marandure et al. (2011) Hubbard BWG,FCR,FI

Chronic Günal (2013) Ross 308 BWG,FCR,FI,Tb Chronic

Yahav and Hurwitz (1996) Cobb BWG,FCR,FI,Tb Acute Bengharbi et al. (2016a)

Hubbard 15 Tb Acute Zulkifli et al. (2003) Cobb Tb

Acute Yalçin et al. (2005) Non- specified Tb Chronic

Abdel-Fattah et al. (2018) Cobb 500 BWG,FCR,FI None

BWG: body weight gain; FI: feed intake; FCR: feed conversion ratio; Tb: body temperature.

Fig. 3. Bubble chart of the design of studies selected for the meta-analysis (n = 17 studies). The x-axis represents the studies included in the meta-analysis. The y-axis indicates the duration of the thermal condi­ tioning applied in each study (when treat­ ments evaluated a range of duration, the average value of all treatments was re­ ported). The diameter of the bubble specifies the temperature of conditioning applied in each study (when treatments evaluated a range of temperature, the average value of all treatments was reported). The color of the bubble designates the category of heat challenge broilers were submitted to in each study.

Table 2 Descriptive statistics of the response variables selected from studies used in the meta-analysis.

N Mean Median Min Max SE BWG (g) 50 1702.2 1890 94.7

2849.5 98 FI (g) 48 30005.8 3462.5 37.7 4940 219.3

FCR 40 1.92 1.82 1.38 2.7 0.05 Tb (◦C) 57 43.2 42.7

40.32 46.37 0.2 BWG: body weight gain; FI: feed intake; FCR: feed conversion ratio; Tb: body temperature.

C.M. Ncho et al.

Journal of Thermal Biology 98 (2021) 102916 5 2001). However, it is critical to make reference and there might be a correlation between the period of conditioning and the extent of sub­ sequent compensatory growth. Several studies (Yahav and Hurwitz,

1996; Yahav and McMurtry, 2001) reported no recovery in the growth performance parameters when the thermal conditioning period was extended above 24 h.

Interestingly, our results demonstrated that broiler’s Tb was signif­ icantly lower under heat stress when they were previously submitted to thermal conditioning. Tb has been generally utilized as a marker for heat stress resistance in broilers (Goel, 2021). Broilers due to the absence of sweat glands are more prone to heat stress which induces an increase in their Tb (Ensminger and Oldfield, 1990). Past examinations exhibited that lower Tb was frequently associated with a significant decline in triiodothyronine levels during exposure to heat stress (Yahav, 1999;

Yahav et al., 1997). Triiodothyronine, the major metabolic hormone plays a crucial role in thermoregulation in broilers and is positively correlated with heat production (Lin et al., 2006). In addition, the overall lower Tb observed in thermally conditioned birds may be an indicator of a lower metabolic rate proposing their ability to cope with high housing temperature (May et al., 1987). Furthermore, the subgroup analysis revealed that thermal conditioning was more effective in reducing broiler’s Tb under acute heat stress than chronic heat stress.

Fig. 4. Forest plot of the standardized mean difference and 95% confidence interval of the effect of thermal conditioning on body weight gain of broilers. The dashed vertical line represents a standardized mean difference of zero, or no effect. Points to the left of the dashed vertical line represent a reduction in body weight gain, while points to the right of the line indicate an increase in body weight gain in broilers that were thermally conditioned.

Fig. 5. Forest plot of the standardized mean difference and 95% confidence interval of the effect of thermal conditioning on feed intake of broilers. The dashed vertical line represents a standardized mean difference of zero, or no effect. Points to the left of the dashed vertical line represent a reduction in feed intake, while points to the right of the line indicate an increase in feed intake in broilers that were thermally conditioned.

C.M. Ncho et al.

Journal of Thermal Biology 98 (2021) 102916 6 The thermotolerance improvement observed after thermal conditioning might be explained by the concept of reactive physiology suggesting a long-lasting memory mechanism (Yahav et al., 1997). Undoubtedly, a large portion of the studies included in this meta-analysis were applying thermal conditioning as short-term acute heat stress at an early age.

Therefore, submitting broilers to the same category of stress in their later life could activate their thermoregulatory memory acquired during their early life.

Through a meta-analysis approach, this study attempted to synthe­ size the literature and provide a quantitative analysis of the effects of thermal conditioning on growth performances and thermotolerance of broilers. However, some limitations exist within the present study.

Firstly, because of the shortage of records, BWG, FI, and FCR were recorded only under standard housing temperature. Indeed, studies that evaluated acute heat stress in their design could not provide the related growth parameters. Therefore, the effects of thermal conditioning on growth performances of broilers under heat stress could not be evalu­ ated. Secondly, significant heterogeneity was present in the response traits, thus it should be taken into consideration when using the result of this study.

5. Conclusion Growth performances are the important parameter for broiler chickens. The meta-analysis highlighted the positive effect of thermal conditioning on broiler growth performances, especially regarding BWG and FI irrespective of the heat stress conditions. Body temperature is a vital indicator of stress during high ambient temperature. Early age

Fig. 6. Forest plot of the mean difference and 95% confidence interval of the effect of thermal conditioning on feed conversion ratio of broilers. The dashed vertical line represents a mean difference of zero, or no effect. Points to the left of the dashed vertical line represent a reduction in feed conversion ratio, while points to the right of the line indicate an increase in feed conversion ratio in broilers that were thermally conditioned.

Fig. 7. Body temperature records extracted from the studies included in the meta-analysis (n = 17 studies). (A) Evolution of broiler’s body temperature based on the heat challenge temperature. (B) Distribution of broiler’s body temperature according to the heat challenge category. Abbreviations: CON: Control; TC: Thermal conditioned.

C.M. Ncho et al.

Journal of Thermal Biology 98 (2021) 102916 7 thermal conditioning effectively reduces the broiler’s Tb under acute heat stress. The effect of thermal conditioning on chronic heat stress broilers was limited in terms of Tb, therefore, other strategies should be considered for helping broilers to cope with chronic heat stress.

Author statement Akshat Goel and Chris Major Ncho: Conceptualization, Methodology.

Chris Major Ncho and Vaishali Gupta: Data curation, Writing- Original draft preparation.Akshat Goel: Supervision. Akshat Goel and Chris

Major Ncho:Writing- Reviewing and Editing.

Fig. 8. Forest plot of the standardized mean difference and 95% confidence interval of the effect of thermal conditioning on body temperature of broilers after heat stress. The dashed vertical line represents a standardized mean difference of zero, or no effect. Points to the left of the dashed vertical line represent a reduction in body temperature, while points to the right of the line indicate an increase in body temperature in broilers that were thermally conditioned.

Fig. 9. Publication bias analysis, funnel plot. Egger’s Linear regression test of funnel plot asymmetry: A) body weight gain (t = 0.06, df = 31, P = 0.9524); B) feed intake (t = -0.40, df = 29, P = 0.6925); C) feed conversion ratio (t = 0.65, df = 24, P = 0.5245); D) body temperature (t = 0.84, df = 30, P = 0.4098).

C.M. Ncho et al.

Journal of Thermal Biology 98 (2021) 102916 8 Declaration of competing interest

The authors declare no conflicts of interest.

References Abdel-Fattah, S., Shourrap, M., Hemida, M.A., 2018. Effect of pre-and post-hatch thermal conditioning on productive performance, some metabolic hormones and breast muscles growth of broilers chicks. Asian J. Anim. Vet. Adv. 13, 369–376.

Abdelqader, A., Al-Fataftah, A.-R., 2014. Thermal acclimation of broiler birds by intermittent heat exposure. J. Therm. Biol. 39, 1–5.

Awad, E.A., Najaa, M., Zulaikha, Z.A., Zulkifli, I., Soleimani, A.F., 2020. Effects of heat stress on growth performance, selected physiological and immunological parameters, caecal microflora, and meat quality in two broiler strains. Asian- Australas. J. Anim. Sci. 33, 778.

Bengharbi, Z., Dahmouni, S., Mouats, A., Petkova, M., Halbouche, M., 2016a.

Physiological variations during a gradual six-hour simulated heat stress in early-age acclimated broilers fed linseed supplemented diet. Bulg. J. Agricult. Sci. 22, 25–33.

Bengharbi, Z., Dahmouni, S., Mouats, M., Halbouche, M., Petkova, M., 2016b. Dietary linseed inclusion and early-age acclimation effects on carcass yield, organs development and thermal resistance of broilers in hot climate. Bulg. J. Agricult. Sci.

22, 34–41.

De Basilio, V., Vilarino, M., Yahav, S., Picard, M., 2001. Early age thermal conditioning and a dual feeding program for male broilers challenged by heat stress. Poultry Sci.

80, 29–36.

De Toledo, T.D.S., Roll, A.A.P., Rutz, F., Dallmann, H.M., Dai Pr´a, M.A., Leite, F.P.L.,

Roll, V.F.B., 2020. An assessment of the impacts of litter treatments on the litter quality and broiler performance: a systematic review and meta-analysis. PloS One

15, e0232853.

DerSimonian, R., Kacker, R., 2007. Random-effects model for meta-analysis of clinical trials: an update. Contemp. Clin. Trials 28, 105–114. https://doi.org/10.1016/j. cct.2006.04.004.

Ensminger, M.E., Oldfield, J.E., 1990. Feeds & Nutrition.

Farag, M.R., Alagawany, M., 2018. Physiological alterations of poultry to the high environmental temperature. J. Therm. Biol. 76, 101–106. https://doi.org/10.1016/j. jtherbio.2018.07.012.

Goel, A., 2021. Heat stress management in poultry. J. Anim. Physiol. Anim. Nutr. https://doi.org/10.1111/jpn.13496.

Günal, M., 2013. The effects of early-age thermal manipulation and daily short-term fasting on performance and body temperatures in broiler exposed to heat stress.

J. Anim. Physiol. Anim. Nutr. 97, 854–860.

Kang, D., Park, J., Shim, K., 2019. Heat treatment at an early age has effects on the resistance to chronic heat stress on broilers. Animals 9, 1022.

Kinney, J.M., 1992. Energy Metabolism: Tissue Determinants and Cellular Corollaries. raven Press.

Liew, P., Zulkifli, I., Hair-Bejo, M., Omar, A., Israf, D., 2003. Effects of early age feed restriction and heat conditioning on heat shock protein 70 expression, resistance to infectious bursal disease, and growth in male broiler chickens subjected to heat stress. Poultry Sci. 82, 1879–1885.

Lin, H., Decuypere, E., Buyse, J., 2006. Acute heat stress induces oxidative stress in broiler chickens. Comp. Biochem. Physiol. Part A Mol. Integr. Physiol. 144, 11–17. https://doi.org/10.1016/j.cbpa.2006.01.032.

Marandure, T., Hamudikuwanda, H., Mashonjowa, E., 2011. Effect of duration of early age thermal conditioning on growth and heat tolerance in broiler chickens. Electron.

J. Environ. Agric. Food Chem. 10.

May, J.D., Deaton, J.W., Branton, S.L., 1987. Body temperature of acclimated broilers during exposure to high temperature. Poultry Sci. 66, 378–380. https://doi.org/

10.3382/ps.0660378.

Modrey, P., Nichelmann, M., 1992. Development of autonomic and behavioural thermoregulation in turkeys (Meleagris gallopavo). J. Therm. Biol. 17, 287–292.

Nawab, A., Ibtisham, F., Li, G., Kieser, B., Wu, J., Liu, W., Zhao, Y., Nawab, Y., Li, K.,

Xiao, M., An, L., 2018. Heat stress in poultry production: mitigation strategies to overcome the future challenges facing the global poultry industry. J. Therm. Biol. 78,

131–139. https://doi.org/10.1016/j.jtherbio.2018.08.010.

Oke, O., Alo, E., Oke, F., Oyebamiji, Y., Ijaiya, M., Odefemi, M., Kazeem, R., Soyode, A.,

Aruwajoye, O., Ojo, R., 2020. Early age thermal manipulation on the performance and physiological response of broiler chickens under hot humid tropical climate.

J. Therm. Biol. 88, 102517.

Plavnik, I., Hurwitz, S., 1985. The performance of broiler chicks during and following a severe feed restriction at an early age. Poultry Sci. 64, 348–355.

Sandercock, D., Hunter, R.R., Mitchell, M., Hocking, P., 2006. Thermoregulatory capacity and muscle membrane integrity are compromised in broilers compared with layers at the same age or body weight. Br. Poultry Sci. 47, 322–329.

Taouis, M., De Basilio, V., Mignon-Grasteau, S., Crochet, S., Bouchot, C., Bigot, K.,

Collin, A., Picard, M., 2002. Early-age thermal conditioning reduces uncoupling protein messenger RNA expression in pectoral muscle of broiler chicks at seven days of age. Poultry Sci. 81, 1640–1643.

Uni, Z., Gal-Garber, O., Geyra, A., Sklan, D., Yahav, S., 2001. Changes in growth and function of chick small intestine epithelium due to early thermal conditioning.

Poultry Sci. 80, 438–445.

Xie, J., Tang, L., Lu, L., Zhang, L., Lin, X., Liu, H.-C., Odle, J., Luo, X., 2015. Effects of acute and chronic heat stress on plasma metabolites, hormones and oxidant status in restrictedly fed broiler breeders. Poultry Sci. 94, 1635–1644.

Yahav, S., 1999. Effect of early-age thermal conditioning and food restriction on performance and thermotolerance of male broiler chickens. Br. Poultry Sci. 40,

120–126.

Yahav, S., Hurwitz, S., 1996. Induction of thermotolerance in male broiler chickens by temperature conditioning at an early age. Poultry Sci. 75, 402–406.

Yahav, S., McMurtry, J., 2001. Thermotolerance acquisition in broiler chickens by temperature conditioning early in life—the effect of timing and ambient temperature. Poultry Sci. 80, 1662–1666.

Yahav, S., Shamai, A., Haberfeld, A., Horev, G., Hurwitz, S., Einat, M., 1997. Induction of thermotolerance in chickens by temperature conditioning: heat shock protein expression. Ann. N. Y. Acad. Sci. 813, 628–636.

Yalçin, S., ¨Ozkan, S., Cabuk, M., Buyse, J., 2005. Pre-and postnatal conditioning induced thermotolerance on body weight, physiological responses and relative asymmetry of broilers originating from young and old breeder flocks. Poultry Sci. 84, 967.

Zaboli, G.-R., Rahimi, S., Shariatmadari, F., Torshizi, M.A.K., Baghbanzadeh, A.,

Mehri, M., 2017. Thermal manipulation during Pre and Post-Hatch on thermotolerance of male broiler chickens exposed to chronic heat stress. Poultry Sci.

96, 478–485.

Zulkifli, I., Liew, P., Israf, D., Omar, A., Hair-Bejo, M., 2003. Effects of early age feed restriction and heat conditioning on heterophil/lymphocyte ratios, heat shock protein 70 expression and body temperature of heat-stressed broiler chickens.

J. Therm. Biol. 28, 217–222.

C.M. Ncho et al.

📖 中文全文 Chinese Full Text

中文

# 热调节对肉鸡生长性能和耐热性的影响:系统综述与荟萃分析

**Chris Major Ncho a, Vaishali Gupta b, Akshat Goel a,\***

a 庆尚国立大学动物科学系,韩国晋州 52828 b 庆尚国立大学应用生命科学系,韩国晋州 52828

---

## 摘要

热调节作为一种经济有效的方法被引入,用于改善肉鸡的生长性能和耐热性。然而,由于各项试验均在不同的实验条件下进行,因此难以得出普遍性结论。因此,本研究旨在通过荟萃分析方法量化肉鸡对热调节的反应。文献检索于2020年12月在Scopus、PubMed、Scielo、Web of Science和Google Scholar上进行。采用限制性最大似然随机效应模型对体增重(BWG)、采食量(FI)、料肉比(FCR)和体温(Tb)的效应量进行合并。BWG、FI和Tb采用标准化均差(SMD)计算,而FCR采用均差(MD)计算,置信区间(CI)为95%。生长性能在热中性条件下进行评估,而Tb在早期热调节后进行急性或慢性热应激后评估。共有17项研究被纳入数据集。与对照组相比,热调节显著提高了BWG(SMD = 0.139,CI = 0.0372–0.2407,P = 0.0074)和FI(SMD = 0.292,CI = 0.108–0.476,P = 0.0019)。此外,亚组分析显示,在急性热应激下Tb显著降低(SMD = -0.455,CI = -0.718至-0.192,P < 0.001),但在慢性热应激下未受影响(SMD = -0.115,CI = -0.651至-0.420,P = 0.6729)。总之,热调节在热中性条件下显著提高了肉鸡的BWG和FI,并有助于在急性热应激下降低Tb。

**关键词:** 热调节;肉鸡;热应激;荟萃分析

---

## 1. 引言

随着全球气候变化,环境变化成为动物生产面临的挑战(Nawab等,2018)。尤其在禽类养殖中,现代肉鸡品种在温和气候条件下已被广泛选育以提高其生长性能(Sandercock等,2006)。此外,汗腺的缺失以及羽毛的存在意味着鸡在高温环境中无法有效调节体温(Xie等,2015)。

多项研究表明,早期热调节可被视为一种减轻肉鸡生产中高温环境不利影响的技术(De Basilio等,2001;Oke等,2020;Yahav,1999)。尽管研究人员尝试了不同的温度和时间周期,但热调节的方法相似,即温度保持在略高于推荐温度的水平,持续数小时甚至数天(图1)。通过热调节提高肉鸡耐热性之所以可行,是因为在其生命的第一周,体温调节机制尚未成熟(Modrey和Nichelmann,1992),这与交感神经活动和下丘脑中的热信息整合密切相关(Kinney,1992)。此外,有证据表明,对肉鸡进行热调节可以促进生长性能的更好发育以及更高的胴体产量(Bengharbi等,2016b;Uni等,2001)。

然而,尽管已有大量研究探索热调节的作用,但文献中获得的结果之间存在诸多分歧。因此,评估先前发表的数据以发现热调节在肉鸡生产中的潜在有效性至关重要。本研究旨在通过荟萃分析方法量化肉鸡对热调节的反应。生长参数如体重、采食量和料肉比在热中性条件下进行评估,而体温在高温条件下进行评估。

---

## 2. 材料与方法

### 2.1. 检索与数据筛选

研究通过查阅以下电子数据库进行:Scopus、PubMed、Scielo、Web of Science和Google Scholar,检索时间为2020年12月。在如此广泛的数据库中进行检索可避免仅纳入单一数据库中的文章。检索关键词包括:"thermal acclimation"、"thermal conditioning"、"heat conditioning"、"temperature conditioning"、"broilers"及其不同组合。两名独立研究人员进行检索。纳入数据库前的研究通过标题、摘要和全文进行严格分析。仅在两名研究人员达成一致后才选择某项研究。未联系作者以获取更多信息或未发表数据。

选择已发表文章的主要标准为:(i)文章应以英文发表在同行评审期刊上;(ii)以肉鸡为实验动物;(iii)文章应至少包含一种孵化后热调节处理;(iv)所有热调节参数(温度、持续时间、起始日龄)应在材料与方法中可获取;(v)文章应评估热调节(冷调节研究未纳入数据库);(vi)对于评估热应激反应的研究,应在热应激条件下设置对照处理。经过筛选步骤后,最终数据库包含从17篇论文中收集的数据。

### 2.2. 数据记录与编码

从研究中提取的响应性状分类如下:生长参数和体温。从每项研究中提取均值、标准差或均值标准误(SEM)以及每个处理的动物数量。当未报告标准差时,根据均值标准误进行计算。由于报告热应激下生长性能的数据不足,且仅考虑热调节的影响,因此仅将标准饲养温度下报告的生长参数纳入数据集。考虑的性状包括体增重(BWG)、采食量(FI)和料肉比(FCR)。热挑战下的体温(Tb)是提取的最后一个响应性状。

### 2.3. 统计分析

使用R软件4.0.3版本(R Core Team,2020)的"metafor"包进行所有统计分析和图形生成。采用限制性最大似然随机效应模型分析数据集。由于响应性状的性质不同,除FCR转换为均差(MD)外,其余均采用标准化均差(SMD)计算,如De Toledo等(2020)先前所执行。关于Tb,通过划分不同热应激类别进行亚组分析,以评估异质性结果并进行组间比较。考虑的两个亚组为慢性和急性热应激。研究根据其所考虑的热应激类别被纳入一个组或另一个组。通过计算卡方检验(Q)和I²统计量评估异质性。若P值<0.05(Q)和/或I² > 50%,则认为存在显著异质性。最后,通过Egger检验和漏斗图评估发表偏倚。

---

## 3. 结果

### 3.1. 研究选择过程

研究选择过程的详细信息如图2所示。对电子数据库中1996年至2020年发表的文章进行文献检索,共检索到8040项研究。去除1348篇重复文章后,剩余6692篇文章。对这些文章的摘要和标题进行严格筛选后,排除了6662篇与我们的主题不直接相关的记录。随后,从保留的30篇文章中,11篇因全文分析后缺乏对研究有帮助的重要数据而被排除。此外,2篇评估冷调节的文章未被选择,因为本综述的目的仅是评估热调节的影响。

### 3.2. 数据集描述

表1显示了入选荟萃分析的文章特征。研究中使用的品系频率如下:Arbor Acres(n = 1);Ross(n = 3);Hubbard(n = 3);Cobb(n = 7),3项研究未指定肉鸡品系。图3展示了入选荟萃分析研究的泡泡图。调节温度差异很大,从35°C到40.5°C不等。大多数研究应用的调节持续时间为1天(n = 11)。共有4项研究在其设计中未包含热挑战,6项选择慢性热应激,7项选择急性热应激。总体热应激温度变化范围为30°C至43°C。报告最多的响应性状是BWG(14项研究),其次是FI(12项研究)、FCR(9项研究)和Tb(9项研究)。所评估响应性状的汇总统计量见表2。

### 3.3. 热调节对肉鸡生长性能的影响

森林图(图4)表明,与对照组相比,热调节对鸟类的BWG有总体正向影响(SMD = 0.139,CI = 0.0372–0.2407,P = 0.0074)。该效应存在异质性,I² = 57.7%,P值<0.001。同样,在图5中,菱形位置反映了热调节对肉鸡FI的正向影响。总体效应估计为SMD = 0.292,CI = 0.108–0.476,P = 0.0019,存在显著异质性(I² = 85.7%,P值<0.001)。图6显示,热调节对肉鸡FCR无显著影响(MD = -0.013,CI = -0.0307–0.0041,P = 0.1346)。此外,观察到显著异质性(I² = 89.7%,P值<0.001)。

### 3.4. 热调节对肉鸡体温的影响

在热挑战期间,在两个极端环境温度(30°C和43°C)下,热调节鸟与对照组之间的体温差异更大,而在36°C至39°C之间,两组的体温相似(图7A)。在急性热应激期间,鸟类的总体体温高于评估慢性热应激的研究收集的数据(图7B)。森林图中的菱形(图8)表明,在热应激下,热调节鸟的Tb总体低于对照组(SMD = -0.315,CI = -0.584至-0.046,P = 0.0219)。该效应具有显著异质性(I² = 73.9%,P值<0.001)。亚组分析显示,在急性热应激期间,热调节鸟的体温显著降低(SMD = -0.455,CI = -0.718至-0.192,P < 0.001)。然而,在慢性热应激下,热调节与对照组相比在Tb方面未显示任何统计学显著差异(SMD = -0.115,CI = -0.651至-0.420,P = 0.6729)。

**发表偏倚。** 图9显示了本荟萃分析中研究参数的漏斗图。所有漏斗图的对称性通过Egger线性回归检验得到确认。BWG(t = 0.06,df = 31,P = 0.9524)、FI(t = -0.40,df = 29,P = 0.6925)、FCR(t = 0.65,df = 24,P = 0.5245)和Tb(t = 0.84,df = 30,P = 0.4098)的结果均不显著。因此,本研究中未检测到发表偏倚风险。

---

## 4. 讨论

畜牧业竞争激烈。广泛的选育和营养规划强调需要识别改善肉鸡生长性能和耐热性的新方法。过去十年中,已进行了多项评估早期热调节对肉鸡生长性能和耐热性影响的试验。然而,总结该主题先前实验结果的最新信息仍然匮乏。因此,本综述试图通过统计荟萃分析方法评估热调节在肉鸡生产中的潜在结果。在本研究中,由于在所有评估的响应性状中检测到中度至高度异质性,因此应用了随机效应模型。事实上,随机效应方法明确考虑了代表研究间变异的异质性(DerSimonian和Kacker,2007)。

当前研究的主要结果之一是,热调节显著提高了肉鸡的BWG和FI,而FCR未受影响。一般来说,热调节期间短期暴露于高温会导致随后的生长迟缓(Yahav和Hurwitz,1996;Yahav等,1997)。据信,热调节肉鸡BWG和FI的最终增加是代偿性生长的结果,类似于早期限饲期间观察到的现象(Plavnik和Hurwitz,1985)。事实上,已经公认高饲养温度会导致肉鸡FI减少,从而导致体重下降(Awad等,2020;Farag和Alagawany,2018)。因此,调节后生长加速以完全补偿损失,有时在上市日龄时导致更高的体重(Yahav和McMurtry,2001)。然而,需要指出的是,调节持续时间与随后的代偿性生长程度之间可能存在相关性。多项研究(Yahav和Hurwitz,1996;Yahav和McMurtry,2001)报告,当热调节持续时间延长至24小时以上时,生长性能参数未见恢复。

有趣的是,我们的结果表明,当肉鸡先前接受过热调节时,热应激下的Tb显著降低。Tb通常被用作肉鸡耐热性的标志物(Goel,2021)。由于缺乏汗腺,肉鸡更容易受到热应激的影响,这会导致其Tb升高(Ensminger和Oldfield,1990)。过去的研究表明,在热应激暴露期间,较低的Tb通常与三碘甲状腺原氨酸水平的显著下降相关(Yahav,1999;Yahav等,1997)。三碘甲状腺原氨酸作为主要的代谢激素,在肉鸡的体温调节中起关键作用,并与产热呈正相关(Lin等,2006)。此外,在热调节鸟类中观察到的总体较低的Tb可能是代谢率降低的指标,表明它们应对高饲养温度的能力(May等,1987)。此外,亚组分析显示,热调节在降低急性热应激下的肉鸡Tb方面比慢性热应激更有效。

热调节后观察到的耐热性改善可以通过反应性生理学概念来解释,该概念表明存在一种持久的记忆机制(Yahav等,1997)。毫无疑问,本荟萃分析中包含的大部分研究在早期将热调节作为短期急性热应激应用。因此,在其生命后期对肉鸡施加相同类别的应激可能会激活其在早期获得的体温调节记忆。

通过荟萃分析方法,本研究试图综合文献并提供热调节对肉鸡生长性能和耐热性影响的定量分析。然而,本研究存在一些局限性。首先,由于记录不足,BWG、FI和FCR仅在标准饲养温度下记录。事实上,在其设计中评估急性热应激的研究无法提供相关的生长参数。因此,无法评估热调节对热应激下肉鸡生长性能的影响。其次,响应性状中存在显著异质性,因此在使用本研究结果时应予以考虑。

---

## 5. 结论

生长性能是肉鸡的重要参数。荟萃分析强调了热调节对肉鸡生长性能的正向影响,特别是在BWG和FI方面,无论热应激条件如何。体温是高温环境下应激的重要指标。早期热调节有效降低了急性热应激下肉鸡的Tb。热调节对慢性热应激肉鸡在Tb方面的影响有限,因此应考虑其他策略来帮助肉鸡应对慢性热应激。

---

**作者声明**

Akshat Goel和Chris Major Ncho:概念化、方法论。Chris Major Ncho和Vaishali Gupta:数据整理、初稿撰写。Akshat Goel:监督。Akshat Goel和Chris Major Ncho:审阅与编辑。

**利益冲突声明**

作者声明无利益冲突。