Introduction
Moulting is an episode of natural rejuvenation in the life of most avian species, including both wild and domestic. Moulting refers to periodic shedding and feather replacement, coinciding with a decrease in the reproductive efficiency function (Berry, Reference BERRY2003) and body weight loss of about 25% (Mrosovsky and Sherry, Reference MROSOVSKY and SHERRY1980). Induced moulting is not new to the poultry industry, as it came into practice in early 20th century (Rice, Reference RICE1905; Rice et al., Reference RICE, NIXON and ROGERS1908). This practice became the most adopted tool to maintain the production potential of laying hens until the early eighties. Most commercial farmers in the egg producing regions of the world adopted this technique until the start of this decade. Due to the widespread criticism of fasting or severe feed-restriction moult induction methods, scientists have developed various alternate methods of moulting. These include high dietary zinc supplementation (Sandhu et al., Reference SANDHU, RAHMAN and RAHMAN2006), low calcium (Breeding et al., Reference BREEDING, BRAKE and GARLICH1992) and low sodium diets (Berry and Brake, Reference BERRY and BRAKE1985), wheat middling (Biggs et al., Reference BIGGS, DOUGLAS, KOELKEBECK and PARSONS2003), using barley (Onbasilar and Erol, Reference ONBASILAR and EROL2007), cottonseed meals (Davis et al., Reference DAVIS, LORDELO and DALE2002), jojoba meal (Vermaut et al., Reference VERMAUT, DE-CONİNCK, FLO, COKELAERE, ONAGBESAN and DECUYPERE1997), alfalfa (Donalson et al., Reference DONALSON, KIM, WOODWARD, HERRERA, KUBENA, NISBET and RICKE2005; Landers et al., Reference LANDERS, WOODWARD, LI, KUBENA, NISBET and RICKE2005; Aygun and Olgun, Reference AYGUN and OLGUN2010), oat (Fletcher et al., Reference FLETCHER, BRİTTON, RAHN and SAVAGE1981; Aygun and Yetisir, Reference AYGUN and YETISIR2009) and rice bran (Santos et al., Reference SANTOS, GARCIA, VIEIRA-FILHO, MOLINO, PELÍCIA, BERTO, MURAKAMI and MONTENEGRO2014) in diets.
The most significant benefit of forced moulting is the restoration of reproductive performance of hens resulting from improved tissue efficiency, loss of adipose tissue, gonadal development and the re-development of tissues and organs. All of these factors ultimately improve post-moult performance (Berry, Reference BERRY2003; Park et al., Reference PARK, KIM, BIRKHOLD, KUBENA, NISBET and RICKE2004). Likewise, it is well known that forced moulting improves male reproductive performance by improving the volume of semen and sperm number per ejaculate (Woodward et al., Reference WOODWARD, OGASAWARA, SNYDER and STINNETT1975; Khan et al., Reference KHAN, JAVED and MUHAMMAD2013b). The phenomenon of moulting of poultry birds has been in practice for many years with great success; however, this success can be extended to even a higher level by providing new ideas to facilitate moulting, which are discussed in the following review.
Post-moult supplementations
Supplementation of poultry feed with beneficial additives has been in practice for a long time. Various feed supplements such as vitamins, minerals, probiotics and prebiotics are commonly used by the poultry industry. However, the use of these supplements in the diet of post moult poultry birds is a new practice. Merging the phenomenon of moulting with the supplementation of beneficial feed additives is believed to increase the advantageous effects of simple moulting. The health status of the moulted birds could be enhanced by providing various supplementation protocols which may be beneficial in reducing the stress of induced moulting as well.
Role of vitamins
Vitamin C (ascorbic acid) is a non-essential vitamin for avian species because they naturally produce the gluconolactone oxidase enzyme that is necessary for the biosynthesis of ascorbic acid (Lin et al., 2006). Therefore, no recommendation for the addition of ascorbic acid in poultry feed has been established in feed standards (NRC, 1994). However, in a study by Zapata and Gernat (Reference ZAPATA and GERNAT1995) on moulted white leghorn hens, a significantly increased egg shell weight and increased egg production in the vitamin C supplemented group was observed. Furthermore, Khan et al. (Reference KHAN, RAHMAN, JAVED and MUHAMMAD2014) observed that vitamin C supplementation in post moult broiler breeders improved the cellular immunity of the birds by increasing macrophage engulfment percentage. In post moult male birds, vitamin C is considered beneficial in improving reproductive performance by significantly reducing the dead sperm concentration and decreasing oxidative stress of seminal plasma thorough enhancing antioxidant defence of post-moult broiler breeders (Khan et al., Reference KHAN, JAVED and MUHAMMAD2013b; Reference KHAN, RAHMAN, JAVED and MUHAMMAD2013c). It is well reported that induction of moulting causes severe oxidative damage (Anwar et al., Reference ANWAR, RAHMAN, JAVED and MUHAMMAD2012b). This stress can constitute a significant factor of physiological damage and is regarded as the cause of several pathological conditions that affect poultry growth and development (Avanzo et al., Reference AVANZO, DE-MENDOCA, PICCOLI and DE-CERQUEIRA2001; Iqbal et al., Reference IQBAL, CAWTHON, WIDEMAN and BOTTJE2001). Vitamin C is considered very helpful in ameliorating this stress by improving the oxidative stress index (OSI) by decreasing total oxidant status (TOS) and increasing total antioxidant capacity (TAC) when supplemented in feed for post moult broiler breeders (Khan et al., Reference KHAN, RAHMAN, JAVED and MUHAMMAD2013a) and layer breeders (Iftikhar et al., Reference IFTIKHAR, KHALIQ, KHAN, RAHMAN, RAHMAN, ANWAR, JAVED, MUZAFFAR and MAHMOOD2015). Vitamin C plays a supportive role in the recovery from liver damage caused by induced moulting by reducing the serum levels of ALT and AST (Khan et al., Reference KHAN, RAHMAN, JAVED and MUHAMMAD2013a).
Vitamin D (1, 25-dihydroxycholecalciferol) is an essential vitamin for the maintenance of immune response, egg production, eggshell formation and calcium homeostasis in poultry (Yannakopoulos and Morris, Reference YANNAKOPOULOS and MORRIS1979; Aslam et al., Reference ASLAM, GARLICH and QURESHI1998; Salvador et al., Reference SALVADOR, FARIA, MAZALLI, ITO, FARIA-FILHO and ARAÚJO2009). It is synthesised from 7-dihydrocholesterol present in the skin cells, and the process is catalysed by UV from sunlight. Generally, commercial poultry are indoors; hence, birds are usually deficient in vitamin D and hence it is added in feed. Likewise, to compensate for the stress incurred by moulting the birds require additional vitamin D supplementation. According to Do Nascimento et al. (Reference DO NASCIMENTO, MURAKAMI, GUERRA, OSPINAS-ROJAS, FERREIRA and and FANHANI2014) adding vitamin D in post moult diets significantly improved production performance and egg quality and improved feed intake and feed conversion ratio of post moult layer birds. Vitamin D is required by the embryo in order to mobilise calcium from the shell, thereby improving hatchability (Narbaitz et al., Reference NARBAITZ, TSANG and GRUNDER1987). Kazemi-Fard et al. (Reference KAZEMI-FARD, KERMANSHAHI, REZAEI and GOLIAN2013) reported increased hatchability in moulted broiler breeder hens when given vitamin D supplementation after moulting. Yolk colour index was changed significantly when given vitamin D supplementation in moulted poultry birds (Park et al., Reference PARK, NAMKUNG, AHN and PAIK2005; Kazemi-Fard et al., Reference KAZEMI-FARD, KERMANSHAHI, REZAEI and GOLIAN2013).
Similar to vitamin C and D, vitamin E is an important component of feed as it is needed for the integrity and optimal function of muscular, reproductive, circulatory, nervous and immune systems in poultry (Leshchinsky and Klasing, Reference LESHCHINSKY and KLASING2001; Rengaraj and Hong, Reference RENGARAJ and HONG2015). Vitamin E is not synthesized naturally in the avian body, hence it is highly recommended to supplement it in feed (Chan et al., Reference CHAN, GERSON and SUBRAMANIAM1998). In conjunction with extensive research on its role in pre moult cycle, its advantage during the post moult period is as significant as that seen during first production cycle. Khan et al. (Reference KHAN, RAHMAN, JAVED and MUHAMMAD2014) and Mahmood (Reference MAHMOOD2014) found that, in post moult birds, vitamin E supplementation improved immune response by significantly increasing macrophage engulfment percentage (Khan et al., Reference KHAN, RAHMAN, JAVED and MUHAMMAD2014). Khan et al. (Reference KHAN, RAHMAN, JAVED and MUHAMMAD2013a) observed that vitamin E caused a reduction in oxidative stress caused by moulting stress. In similar research it was found that vitamin E also decreases ALT and AST levels significantly, which are an indicator of liver damage. Khan et al. (Reference KHAN, JAVED and MUHAMMAD2013b) results of semen quality parameters revealed that volume, motility and capacity for fertilising eggs were significantly higher in the vitamin E fed group of post moult male broiler breeders. Vitamin E causes a significant increase in the size and area of FSH, LH gonadotrophs and lactotrophs. These results showed that vitamin E alone may play some role in the enhancement of semen quality and growth of gonadotrophs and lactotrophs. Vitamin E helps improve the metabolic hormones level including T3 and T4 in post moult layer breeders (Iftikhar et al., Reference IFTIKHAR, KHALIQ, KHAN, RAHMAN, RAHMAN, ANWAR, JAVED, MUZAFFAR and MAHMOOD2015) and broiler breeders (Khan et al., Reference KHAN, RAHMAN, JAVED and MUHAMMAD2013a).
Role of probiotics
Probiotics are the mixed culture of living microorganisms, which have a beneficial effect on the host by improving the populations of the native microflora (Houndonougbo et al., Reference HOUNDONOUGBO, CHRYSOSTOME and AMOUSSA2011; Dibaji et al., Reference DIBAJI, SEIDAVI and ASADPOUR2012). The mechanism of action of probiotics includes; microbial antagonism (Fuller, Reference FULLER1989; Kelly and King, Reference KELLY and KING2001; Walsh et al., Reference WALSH, PEDDIREDDI and RADCLIFFE2004; Mountzouris et al., Reference MOUNTZOURIS, BENEAS, TSIRTSIKOS, KALAMARA and FEGEROS2006), competitive exclusion (Alexopoulos et al., Reference ALEXOPOULOS, GEORGOULAKIS, TZIVARA, KRITAS, SIOCHU and KYRIAKIS2004; Berchieri et al., Reference BERCHIERI, STERZO, PAIVA, LUCKSTADT and BELTRAN2006), vitamins synthesis (Coates and Fuller, Reference COATES and FULLER1977), utilising indigestible carbohydrate (Prins, Reference PRINS1977) and immune modulation (Perdigon et al., Reference PERDIGON, FULLER and RAYA2001; Toms and Powrie, Reference TOMS and POWRIE2001; Lan et al., Reference LAN, VERSTEGEN, TAMINGA and WILLIAMS2005).
Similar to its beneficial effects during growing and production in poultry, it is equally useful during the post-moult production period. In a study by Khajali et al. (Reference KHAJALI, KARIMI and QUJEQ2008) when post-moult laying hens were provided with probiotics they had improved egg production and better feed conversion ratio, which was attributed to recovery from moulting induced stress. In moulted hens supplemented with probiotics, faster restoration of egg production was observed by Anwar et al. (Reference ANWAR, RAHMAN, JAVED and MUHAMMAD2012a) in the post-moult production period. In a more recent study, better egg production in probiotic supplemented moulted laying hens has been attributed to the increased size of gonadotrophs in supplemented birds (Anwar and Rahman, Reference ANWAR and RAHMAN2015b). Similarly, Abdelqader et al. (Reference ABDELQADER, IRSHAID and AL-FATAFTAH2013) observed an increase in egg weight, production, mass, eggshell thickness and weight in probiotic-supplemented laying hens during late phase of production when performance was dropping. Increased total antioxidant capacity and lowering total oxidative status and homocysteine concentrations in post-moult layers (Anwar et al., Reference ANWAR, RAHMAN, JAVED and MUHAMMAD2012b) and male layer breeders (Iftikhar et al., Reference IFTIKHAR, KHALIQ, KHAN, RAHMAN, RAHMAN, ANWAR, JAVED, MUZAFFAR and MAHMOOD2015) have been observed in response to probiotic supplementation. Likewise, Sohail et al. (Reference SOHAIL, IJAZ, YOUNUS, SHABBIR, KAMRAN, AHMAD, ANWAR, YOUSAF, ASHRAF, SHAHZAD and REHMAN2013) found that probiotics are helpful in ameliorating heat induced oxidative stress by improving oxidative status of body.
The induction of moulting impairs the cellular and humoral immune response of bird including an increase in heterophil to lymphocyte ratio which is an important indicator of immune stress (Holt and Porter, Reference HOLT and PORTER1992; Siegel, Reference SIEGEL1995). Post-moult probiotic supplementation aids in the reduction of increased heterophil to lymphocyte ratio and is thus considered helpful in reducing immune stress incurred by moulting (Khajali et al., Reference KHAJALI, KARIMI and QUJEQ2008). Khan et al. (Reference KHAN, JAVED and MUHAMMAD2013b) and Mahmood (Reference MAHMOOD2014) observed improvements in different immune parameters when probiotics were given to post moult male broiler breeders. Anwar et al. (Reference ANWAR, RAHMAN, JAVED and MUHAMMAD2015a) reported an improvement in humoral immunity of moulted white leghorn hens given probiotics. Hence, overall it can easily be concluded that probiotics supplementation during the post-moult period is helpful in revoking the oxidative and immune stress caused by induced moulting.
Role of protein
Crude protein (CP) is an important parameter to modulate growth, reproductive performance and development in poultry (Forbes and Shariatmadari, Reference FORBES and SHARIATMADARI1994). Similar to its importance during the pre-moult period, it is equally essential for post-moult birds.
In a study by Anwar et al. (Reference ANWAR, RAHMAN, JAVED and MUHAMMAD2012a), immunohistochemistry of the pituitary gland revealed a significant decrease in size and area of lactotrophs in post-moult layers fed a high protein diet. This indicated earlier post-moult recovery with better egg production in the protein supplemented group as compared to control group. In another study by Iftikhar et al. (Reference IFTIKHAR, KHALIQ, KHAN, RAHMAN, RAHMAN, ANWAR, JAVED, MUZAFFAR and MAHMOOD2015), when post-moult layer birds were given high protein supplementation, a significant increase in total antioxidant capacity (TAC), paraoxonase, ceruloplasmin and arylesterase activity while a significant decrease in total oxidant status (TOS) and homocysteine levels were observed, which indicated a significant decrease in oxidative stress caused by moulting. Similar results were found by Khan et al. (Reference KHAN, JAVED and MUHAMMAD2013b) in post-moult broiler breeders.
During the post-moult stage, various feeding regimens varying from low to high CP are in practice. Traditional diets are no longer sufficient to provide the protein requirement of post-moult birds; hence, the supplementation with additional protein gives better production results which are of great importance in the moulted layers.
Togun et al. (Reference TOGUN, OKWUSIDI, AMAO and ONYIAOHA2004) studied the effects of varying levels of crude protein and follicle stimulation on the recovery and performance of 100 post-moult Nera Black hens. Follicle stimulation was enhanced in post-moult production in birds fed the 16% CP diet. Koelkebeck et al. (Reference KOELKEBECK, PARSONS, LEEPER and MOSHTAGHIAN1991) observed significantly higher body weight, feed consumption and feed conversion ratio in post-moult laying hens fed a 16.5% CP diet. Hens fed higher protein diets returned to egg production faster than those fed lower protein diets. Javaid et al. (Reference JAVAID, ANJUM and AKRAM2012) recommended that a dietary level of 16% CP may be suitable for hens during the moulting process and increases above this level (up to 18%) leads to wastage of protein. Gunawardana et al. (Reference GUNAWARDANA, ROLAND and BRYANT2008) mentioned that increasing the dietary protein by 2-3% could result in an increase of 6% in feed intake of moulted birds, giving improved egg quality and production.
Among the different amino acids, methionine is one of the most important amino acids and layer diets are generally deficient in this amino acid. Even diets with low CP but supplemented with necessary amino acids, especially methionine (Koelkebeck et al., Reference KOELKEBECK, PARSONS, LEEPER and MOSHTAGHIAN1991; Bunchasak and Silapasorn, Reference BUNCHASAK and SILAPASORN2005) and L-carnitine (Arslan, Reference ARSLAN2006), are adequate to alleviate the problems of protein insufficiency. Wu et al. (Reference WU, BRYANT, VOITLE and ROLAND2005) reported a significant increase in egg weight of moulted hens by increasing the protein, fat, linoleic acid, lysine and methionine. Domingues et al. (Reference DOMINGUES, SGAVIOLI, PRAES, DUARTE, CASTIBLANCO, SANTOS, ALVA and JUNQUEIRA2012) found a significant increase in egg production in post-moult hens when supplemented with lysine and methionine.
CP level is crucial for achieving and maintaining reproductive performance of both male and female poultry birds (Zhang et al., Reference ZHANG, BERRY, MCDANIEL, ROLAND, LIU, CALVERT and WILHITE1999; Laudadio et al., Reference LAUDADIO, PASSANTINO, PERILLO, LOPRESTI, PASSANTINO, KHAN and TUFARELLI2012). Although feeding a high protein diet is considered beneficial for female reproductive performance, both high and low levels of protein in the diet can adversely affect semen quality in male birds (Wilson et al., Reference WILSON, ROWLAND and HARMS1971; Hocking and Bernard, Reference HOCKING and BERNARD1997). To support this, Romero-Sanchez et al. (Reference ROMERO-SANCHEZ, PLUMSTEAD, LEKSRISOMPONG, BRANNAN and BRAKE2008) observed improved fertility rates when fed normal CP diet rather than a high protein diet. A study by Khan et al. (Reference KHAN, JAVED and MUHAMMAD2013b) showed improved semen volume, sperm concentration and motility in post-moult male broiler breeders when given a 14% CP diet. Muzaffar et al. (Reference MUZAFFAR, KHALIQ, KHAN, RAHMAN, MAHMOOD, IFTIKHAR, RAHMAN and MAHMOOD2016) observed significant improvements in the reproductive performance of post-moult male layer breeders supplemented with a 12% protein diet. Although high protein diets are beneficial for improving many health parameters of post-moult poultry, in post-moult male birds a medium range protein diet is recommended instead of a very high protein diet.
Role of minerals
Although minerals are already used in poultry feed (Parker, Reference PARKER2013; Świątkiewicz et al., Reference ŚWIĄTKIEWICZ, ARCZEWSKA-WŁOSEK and JOZEFIAK2014), their use in post-moult feed is a relatively new practice. Khan et al. (Reference KHAN, REHMAN and ANSARI2011) observed a significantly higher egg production when zinc, copper and aluminium were supplemented in diets for post-moult layers. Likewise, Stanley et al. (Reference STANLEY, SHANKLYN, DALEY, GRAY, VAUGHAN, HINTON and HUME2012) recorded greater feed utilisation and egg production in selenium and zinc supplemented groups of post-moult hens. Sirirat et al. (Reference SIRIRAT, LU, HUNG and LIEN2013) observed significantly improved productive and reproductive outcomes with chromium supplementation in post-moult layers. Do Nascimento et al. (Reference DO NASCIMENTO, MURAKAMI, GUERRA, OSPINAS-ROJAS, FERREIRA and and FANHANI2014) found improved egg production rate, better feed conversion and egg weights when calcium was added to layer feed in the second cycle. Hence producers should be encouraged to use these minerals in diets for post-moult poultry to get better subsequent results.
Conclusions
The data available in the literature comprehensively advocates the use of vitamin supplementation for post-moult poultry, especially vitamin C and E, to get maximum performance and recovery from moulting. It is evident from research that induced moulting augments physiological stress, and oxidative stress levels increase in moulted birds. The increased oxidative level of the body directly influences other health parameters. Due to the severity of the impact of different moulting techniques on the welfare, animal care agencies all around the world have focussed on this issue. Keeping in mind the economic benefits of moulting, scientists have started experimenting with novel techniques to minimise the harm produced by free radicals. In addition to other novel techniques, diet supplementation, particularly with vitamins E and C can assist the early recovery from oxidative stress after induced moulting. Other published studies provide good support to endorse the use of probiotics and minerals in the feed. Using various feeding strategies can assist in post moulting recovery for birds, helping with welfare issues as well as supporting second cycle productive performance.