Wooden-Breast, White Striping, and Spaghetti

Meat: Causes, Consequences and Consumer

Perception of Emerging Broiler Meat
Abnormalities
M. Petracci, F. Soglia, M. Madruga, L. Carvalho, Elza Ida, and M. Estévez

Abstract: Ten years ago, the occurrence of macroscopic defects in breasts muscles from fast-growing broilers challenged
producers and animal scientists to label and characterize myopathies wholly unknown. The distinctive white striations
in breasts affected by white striping disorder, the presence of out-bulging and pale areas of hardened consistency in
the so-called wooden breast, and the separation of the fiber bundles in breasts labelled as spaghetti meat, made these
myopathies easily identified in chicken carcasses. Yet, the high incidence of these myopathies and the increasing concern
by producers and retailers led to an unprecedented flood of questions on the causes and consequences of these abnormal
chicken breasts. This review comprehensively collects the most relevant information from studies aimed to understand the
pathological mechanisms of these myopathies, their physicochemical and histological characterization and their impact on
meat quality and consumer’s preferences. Today, it is known that the occurrence is linked to fast-growth rates of the birds
and their large breast muscles. The muscle hypertrophy along with an unbalanced growth of supportive connective tissue
leads to a compromised blood supply and hypoxia. The occurrence of oxidative stress and mitochondrial dysfunction
leads to lipidosis, fibrosis, and overall myodegeneration. Along with the altered appearance, breast muscles affected by
the myopathies display poor technological properties, impaired texture properties, and reduced nutritional value. As
consumer’s awareness on the occurrence of these abnormalities and the concerns on animal welfare arise, efforts are
made to inhibit the onset of the myopathies or alleviate the severity of the symptoms. The lack of fully effective dietary
strategies leads scientists to propose whether “slow” production systems may alternatively provide with poultry meat free
of these myopathies.
Keywords: animal welfare, chicken quality, oxidative stress, spaghetti meat, white striping, wooden breast

Introduction with the faded image of red meats, identified as “probably carcino-
The past two decades have witnessed an increase of consumer genic to humans” by the IARC (IARC, 2015). Last but not least,
preference for chicken meat over other types of muscle foods. The poultry production and consumption are worldwide phenomena
worldwide increased consumption of chicken can be attributed as chicken meat complies with most cultural and religious prin-
to its relative low-cost, the diversity and ease of meat prepara- ciples. In order to meet the growing demand and, in parallel,
tion (Wideman, O’Bryan, & Crandall, 2016), appreciated sensory to optimize broiler production and increase profits, chickens have
properties (Petracci, Mudalal, Soglia, & Cavani, 2015), reported been selected for fast growth and high yields (Petracci et al., 2015).
nutritional and dietary properties (Estévez, 2015), and the recog- The studies carried out by Havenstein, Ferket, and Qureshi (2003)
nized positive image of white meats (of being healthy) compared and Havenstein, Ferket, Grimes, Qureshi, and Nestor (2007) have
soundly illustrated to which extent manipulation of genetics and
feeds, has been able to push the biological boundaries in animal
CRF3-2018-0221 Submitted 9/24/2018, Accepted 1/14/2019. Authors Petracci production over the last decades. The modern Ross 308 broiler
and Soglia are with Dept. of Agricultural and Food Sciences, Alma Mater Studiorum, fed on 2001 feeds, would reach 1.8 kg of body weight (BW) at
Univ. of Bologna, Piazza Goidanich 60, 47521, Cesena, Italy. Authors Madruga and
Carvalho are with Postgraduate program in Food Science and Technology. Dept. of Food
32 days of age with a feed conversion (FC) of 1.47, whereas the
Engineering, Federal Univ. of Paraiba, João Pessoa, Paraı́ba, Brazil. Author Ida is with Athens-Canadian Randombred Control (ACRBC) Strain fed on
Dept. of Food Technology, Londrina State Univ., Londrina, Brazil. Author Estévez 1957 feeds, would not have reached that BW until 101 days of age
is with Meat and Meat Products Research Inst., TECAL Research Group, Univ. with a FC of 4.42 (Havenstein et al., 2003). Irrespective of the
of Extremadura, Avda. Universidad s/n, 10003, Cáceres, Spain. Direct inquiries to feeds, a Ross 308 broiler presents an average BW of approximately
author Estévez (E-mail: [email protected]).
2.4 kg (42 days) while that of ACRBC would be approximately


C 2019 Institute of Food Technologists®

doi: 10.1111/1541-4337.12431 Vol. 00, 2019 r Comprehensive Reviews in Food Science and Food Safety 1
Emerging broiler meat abnormalities . . .

Figure 1–Classification of myopathies in chicken breast. A–Normal breast (without white striations or harden areas and hemorrhages);
B–WS-moderate-2 breast (with striation < 1mm covering the breast surface extensively); C–WS-severe-3 breast (with striation > 1mm covering the
breast surface extensively); D–WS-moderate-2 thigh (with striation <1 mm covering the thigh surface extensively); E–WB-moderate breast (with
focal, hardened, and pale areas, without hemorrhages); F–WB—extremely severe breast (with diffused, hardened and pale areas, without
hemorrhages); G–WB-extremely severe breast (with diffused, hardened and pale areas and hemorrhages); H–simultaneous occurrence of WS and WB
(WS/WB) breast (diffused, hardened, and pale areas and superficial white striations in the cranial part); I–SM-extremely severe breast (tendency
towards separation of the fiber bundles composing the muscle tissue itself).

0.55 kg at the same age (Havenstein et al., 2003). Subsequent in turn, may also appear together with an additional and recently
technical guides published by Aviagen confirm the trend of in- reported myopathy called SM. SM is characterized by an overall
creasing chicken breast yields in Ross 308 broiler (Aviagen, 2007, impaired integrity of the pectoralis major muscle that, exhibiting
2012, 2014). Comparable numbers were subsequently shown by the tendency toward separation of the fiber bundles composing the
the same authors when comparing 1966- compared with 2003- muscle tissue itself, provides an image resembling the long, thin,
type turkeys (Havenstein et al., 2007). At the present time, those solid, and cylindrical appearance of the popular pasta (Baldi et al.,
extraordinary achievements seem to have caused the arise of spon- 2018). WS and WB can be classified into several grades depending
taneous myopathies in broilers and turkeys (Petracci et al., 2015) on the severity of the symptoms (Figure 1).
as a reflection of animal biology fighting back the human in- While the information on the incidence of these myopathies is
tervention. While physiological limits could have been reached limited and, at times contradictory, it is assumed chicken breasts
already, ethical issues may have been surpassed long before as these with abnormalities appear in all countries where fast-growing hy-
emerging myopathies, namely white striping (WS), wooden breast brids are used and that the number is higher than that the chicken
(WB), and spaghetti meat (SM), are closely associated with inten- industry would admit. WS, the most common of the myopathies
sive and exhausting animal production systems (Mutryn, Brannick, under examination in the present review, affects to an overall level
Fu, Lee, & Abasht, 2015; Petracci et al., 2015). of 50% of chicken breasts in Italy, France, Spain, and Brazil (Al-
WS is easily recognized by the occurrence of white striations nahhas et al., 2016; Lorenzi, Mudalal, Cavani, & Petracci, 2014;
following the same direction of the muscle fibers in poultry breast Russo et al., 2015; Carvalho, unpublished data) with those display-
(Figure 1). A microscopic examination of these white stripes re- ing severe degree being around 20% to 30% of the total affected
veals accumulation of lipids and proliferation of connective tissue muscles. In Northeastern Brazil, the share of breasts with WB is
(Kuttappan et al., 2013a). WB myopathy, affecting the pectoralis reported to be between 10% and 20% with a high proportion of
major and occasionally the pectoralis minor in broilers, appears as those breasts concurrently exhibiting WS (Carvalho, unpublished
a focal lesion at approximately 2 weeks of age and subsequently de- data). In Italy, a survey carried out between 2017 and 2018 on
velops as a widespread fibrotic injury (Papah, Brannick, Schmidt, 16,000 breasts identified 42% of the samples being moderate WB
& Abasht, 2017). The first thorough pathological description of and 18% being affected to a severe extent (Petracci, unpublished
WB was carried out by Sihvo, Immonen, and Puolanne (2014). data). The same authors identified around 20% of the samples with
These authors reported a hardened consistency and pale appear- SM defect. The assessment of breasts from birds at 9 weeks of age
ance in the pectoral muscles which is microscopically characterized in the United States, revealed that more than 98% were found to
by polyphasic myodegenerations with fibrosis in the chronic phase. display signs of WS (more than 55% classified as moderate and
Both the aforementioned abnormalities have been reported to co- severe cases) and that around 85% of the samples displayed WB
incide in the same muscle although the existence of concurring (more than 42% were severe or very severe) (Kuttappan, Owens,
pathological underlying mechanisms responsible for their occur- Coon, Hargis, & Vazquez-Anon, 2017). It should be emphasized
rence is a disputable topic (Cruz et al., 2016; Kuttappan, Hargis, & that classification criteria may greatly vary among surveys, so ab-
Owens, 2016; Livingston, Landon, Barnes, & Brake, 2018). WS, solute levels should be taken with care.

2 Comprehensive Reviews in Food Science and Food Safety r Vol. 00, 2019 
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Emerging broiler meat abnormalities . . .

Selecon for increased growth rate

and breast muscle development
Aempt to increase the blood flow
Muscle hypertrophy through ↑ NO (vasodilator)

Compromised blood supply to the fibers Impaired metabolic waste product Phlebis and perivascular
likely exacerbated by: displacement lipid infiltraon
↓ blood vessel density
Accumulaon of compounds with osmoc
properes (i.e. taurine, alanine) leading to Oxidave stress
ssue swelling [↓ anserine and carnosine]
[↑ taurine]

Hypoxia

Inflammaon Mitochondrial
dysfuncon

Myodegeneraon
↑ ∅ of sarcoplasmic reculum Altered nucleode Altered glucose
metabolism metabolism
[↑ purine catabolism]
Impaired calcium homeostasis
Collagen, proteoglycan and
Acvaon of proteases glycosaminoglycan synthesis
and lipases
Regeneraon failure
Regeneraon Fiber necrosis Fibrosis and lipidosis

Figure 2–Schematic representation of the possible etiologies and mechanisms leading to the development of white striping (WS), wooden breast (WB),
and spaghetti meat (SM) abnormalities.

These myopathies have been reported to be linked to rapid 1996). Consistently, recent studies carried out in modern broiler
muscle growth, insufficient vascularization, and oxidative stress strains selected for rapid growth and muscle yield revealed that the
that may lead to tissue degeneration (Kuttappan, Brewer, Apple, pectoralis major muscles were entirely composed of type IIB fibers
Waldroup, & Owens 2012a; Soglia et al., 2016a; Papah, Brannick, and the authors hypothesized whether this condition may increase
Schmidt, & Abasht, 2018; Sihvo et al., 2018). Yet, and despite of their susceptibility to the development of muscular abnormalities
the considerable efforts that have been made in the last years, the (Petracci et al., 2017). In this context, an overall impairment in
precise etiology of these abnormalities remained unclear. Regret- muscle regeneration in fast-growing broilers was recently demon-
tably, the remarkable negative impact of these muscular abnormal- strated to be a consequence of a reduction in satellite cells number
ities on the appearance, technological and nutritional quality, and and in their altered abilities to proliferate and differentiate (Clark
consumer acceptance of breast meat, have been clearly underlined. & Velleman, 2016; Daughtry et al., 2018). In addition, it was re-
Hence, a profound understanding on the underlying causes and the cently found a strong genetic determinism of the WS condition in
search for solutions are needed (Baldi et al., 2018; Mutryn et al., fast-growing chickens (h2 = 0.65; Alnahhas et al., 2016), even if
2015; Soglia et al., 2016a). This review compiles all relevant infor- Bailey, Watson, Bilgili, and Avendano (2015) previously reported
mation in relation to the characteristics, potential etiology, quality lower heritability levels in two commercial pure lines of broiler
traits, and consumer behavior toward these new myopathies and chickens selected for high (h2 = 0.34) or moderate (h2 = 0.18)
provides grounds for future challenges aimed to alleviation. breast meat yield. Alnahhas et al. (2016) pointed also out that WS
is genetically more highly related to the development of pectoralis
Underlying Causes of WB, WS, and SM major muscle (rg = +0.73) than to the overall growth of the body
In the past 50 years, in order to fulfil the worldwide increas- (rg = +0.33). Thus, being the major contributor to the changes in
ing demand for poultry meat, selection programs have been car- breast muscle development and yield, selection might be reason-
ried out to improve the production traits of broiler chickens and ably considered as the main underpinning factor responsible for
develop high growth-rate and breast-yield hybrids. As a conse- the development of these muscular abnormalities. In agreement
quence, outstanding results in broilers’ growth performances and with that, a model was recently developed to determine the con-
body composition were achieved and the number of rearing days tribution of different growth parameters (that is, pectoralis major
necessary to obtain market weight birds was reduced to one-half yield, length, depth) in predicting the severity of the myopathic
(Petracci, Soglia, & Berri, 2017; Tallentire, Leinonen, & Kyriaza- lesions associated with WS and WB. The findings demonstrated
kis, 2018). In spite of that upgraded production profitability, these that all those physical measurements inherent to genetic selection
selection practices profoundly altered muscle architecture (that for fast-growing and high-breast yield hybrids were strictly re-
is, increase muscle fiber diameter and length, increase myofiber lated to the occurrence and the severity of muscular abnormalities
number, reduction in capillary density, and capillary to fiber ratio) (Griffin, Moraes, Wick, & Lilburn, 2018).
and metabolism leading to a shift toward the glycolytic pathway However, the studies carried out in order to quantify genes
(Hoving-Bolin, Kranen, Klont, Gerritsen, & De Greef, 2000). expression and identify putative causative genes leading to the de-
Chicken breast muscles are known to be essentially glycolytic in velopment of WS (Pampouille et al., 2018) and/or WB (Abasht,
function and nature (Papinaho, Ruusunen, Suuronen &, Fletcher, Mutryn, Michalek, & Lee, 2016; Huber, Williams, & Athrey,


C 2019 Institute of Food Technologists® Vol. 00, 2019 r Comprehensive Reviews in Food Science and Food Safety 3
Emerging broiler meat abnormalities . . .

2018; Mutryn et al., 2015; Papah et al., 2018; Zambonelli et al., (Boerboom et al., 2018). Moreover, dilation of the sarcoplasmic
2016) evidenced a complex etiology and a polygenic inheritance reticulum (increased diameter), mitochondrial swelling, vacuo-
of the defect. Indeed, several genes and metabolites were differ- lation, cristae loss, and mitochondrial hyperplasia (identified
entially expressed between the pectoralis major affected by WS as early ultrastructural changes associated with the occurrence
and/or WB and the unaffected (or only mildly affected) cases. of WB) are induced by the osmotic imbalances resulting from
However, since it was not possible to identify biomarkers able to hypoxia and myodegeneration (Sihvo et al., 2018). In addition,
discriminate WS from WB, a common etiology leading to the impaired calcium homeostasis (Boerboom et al., 2018; Mutryn
development of these muscular abnormalities might be reason- et al., 2015; Zambonelli et al., 2016) may lead to the activation of
ably hypothesized. Nevertheless, WS, WB, and SM abnormalities proteases and lipases (Zambonelli et al., 2016) thus contributing to
exhibited a distinctive phenotype that might be considered as an myodegeneration and protein degradation (Petracci et al., 2015).
individual-response to the profound alteration in muscle tissue Then, complex biological reactions and regenerative processes,
induced by genetic selection. aimed at alleviating inflammation and limiting cellular apoptosis
In this context, in the past few years, several studies were and tissue necrosis, take place (Petracci et al., 2017). When my-
carried out in order to identify the underlying mechanisms and odegeneration overtake the regenerative capacity of the muscle,
metabolic pathways involved in the occurrence of WS, WB, and altered nucleotide (as evidence by an increased purine catabolism),
SM muscular abnormalities (Abasht et al., 2016; Boerboom, van and carbohydrate metabolisms are observed (Abasht et al., 2016;
Kempen, Navarro-Villa, & Pérez-Bonilla, 2018; Hubert et al., Papah et al., 2018; Zambonelli et al., 2016) and ultimately result
2018; Mutryn et al., 2015; Papah et al., 2017; Papah et al., in fibrosis and lipidosis, distinctive microscopic traits associated
2018; Sihvo, Airas, Lindén, & Puolanne 2018; Zambonelli et al., with the occurrence of these growth-related abnormalities.
2016). However, the exact etiology and the chronology of events
leading to the development of these defects are only partially
understood and fragmentarily described. Since these muscular Physicochemical and Histological Properties of WB,
abnormalities exhibit similar histological features, a common WS, and SM
underlying mechanism responsible for their occurrence might Classification of myopathies
be hypothesized. A schematic representation of the underlying The identification of myopathies in poultry breasts and estab-
mechanisms leading to the progression of WS, WB, and SM lishing the degree of severity, are carried out by visual examination
abnormalities is shown in Figure 2. An early version of this and/or palpation of chicken breast muscle (Figure 2) (Kuttappan
illustration, reported in a previous review (Petracci et al., 2017), et al., 2013a,b; Mutryn et al., 2015). Breasts affected by WS have
has been updated with findings from recent studies (Boerboom been characterized by the presence of white striations parallel to
et al., 2018; Papah et al., 2017, 2018; Sihvo et al., 2018). muscle fibers. In the breast muscle, stripes usually appear in the
Selection for fast-growing and high-breast development in cranial part of the fillet near the wing attachment and may, or
modern broiler hybrids, achieved through fibers’ hypertrophy, may not, extend along the muscle to the caudal region (Ferreira,
likely resulted in compromised blood and oxygen supply to the Casagrande, Vieira, Dreimeier, & Kindlein, 2014). Typically,
muscle tissue leading to the development of hypoxia (Hoving- breasts with WS are categorized into two grades: moderate and
Bolink et al., 2000; Sihvo, Airas, Lindén, & Puolanne, 2018). This severe. Those with a moderate degree predominantly present
condition might even be exacerbated by the physiology of the striations with a thickness of less than 1 mm, while the severely
pectoralis major muscle: its impressive development (with partic- affected cases, most often, have striations with a thickness greater
ular reference to thickness) might compress the pectoral artery than 1 mm; both easily visible (Kuttappan et al., 2012a,c;
thus further reducing oxygenation and nutrients transportation to Kuttappan, Brewer, Apple, Waldroup, and Owens, 2012b). In
the muscle. This hypothesis is further supported by the evidence addition, Bailey et al. (2015), scored WS using a three-point scale
that the severity of the histological lesions, associated with the oc- of severity: 1 (mild–focal appearance of stripes covering part of
currence of WS, WB, and SM, gradually decreases moving from the breast), 2 (moderate—stripes extensively covering the breast
the skin-facing surface towards the inner section of the pectoralis surface), 3 (severe—very thick stripes with extensive coverage
major (Baldi et al., 2018; Clark & Velleman, 2016; Soglia et al., over the breast surface). This condition, although frequently
2016a). In addition, hypoxic conditions seem to be exacerbated by reported in the breast meat (pectoralis major muscle), can also
the reduced blood vessel density observed in WB affected muscles be seen in the thigh (iliotibialis muscle), tenders (pectoralis minor
(Sihvo et al., 2018) as well as by tissue swelling likely due to the muscle), and drumsticks (gastrocnemius muscle) (Kuttappan et al.,
accumulation of compounds having osmotic properties (that is, 2013c; Zimermann et al., 2012).
taurine and alanine) (Boerboom et al., 2018). WB is characterized by a hardening of the breast muscle, which
Concurrently, it has been found that metabolic waste products may have on its surface paler color, surface hemorrhaging and
displacement might be impaired and associated to the develop- exudate (Kuttappan et al., 2016; Sihvo et al., 2014). This rigidity
ment of phlebitis and perivascular lipid infiltration (Papah et al., can be localized (the muscle presents focal rigid consistency, while
2017; Sihvo et al., 2017). Besides the veins, evidences of vascular other areas have normal consistency) or diffuse (extend through-
pathology affecting the arteries, depicted by arteriosclerosis out the breast meat) (Sihvo et al., 2018). Four categories can be
and atherosclerosis, has been observed by analyzing the genes attributed to WB (Papah et al., 2017; Petracci et al., 2017; Sihvo
differentially expressed between 3-week-old unaffected birds et al., 2017): mild (focally diffused and light firmness), moder-
and WB cases (Papah et al., 2018). Within this context, it has ate (focally diffused with extensive firmness of the breast), severe
been speculated that muscle tissue tries to overcome hypoxia (>75% of the breast being extremely firm and with diffuse cover-
by increasing blood flow through the synthesis of nitric oxide age) and extremely severe (firm breast). Although chicken breasts
that, however, might exacerbate and accelerate the development may present only symptoms from this myopathy, it is often re-
of oxidative stress. Such pro-oxidative environment would ulti- ported to be associated with WS (Kuttappan et al., 2017; Soglia
mately contribute to tissue inflammation and myodegeneration et al., 2016a). It is important to highlight that, although WB is

4 Comprehensive Reviews in Food Science and Food Safety r Vol. 00, 2019 
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Emerging broiler meat abnormalities . . .

commonly detected by manual palpation of the breast, this method (2016), and Mudalal et al. (2015) found similar results for breasts
may not be fully effective. Velleman, Clark, and Tonniges (2017) with combined WS and WB myopathies; while Baldi et al. (2018)
hypothesized that palpable hardness of the WB breast may be a re- reported a higher average weight of breast fillet with SM and si-
sult of the fibrosis resulting from the accumulation of cross-linked multaneous occurrence of WS/WB and WS/SM when compared
collagen fibrils. Other authors reported consistent results and indi- to normal one. In relation to the height of the pectoralis major
cated that collagen may play a major role in defining the increased muscle, chicken breasts affected by WS, WB, SM, WS/WB, and
firmness associated to the development of this condition (Soglia WS/SM had higher height in both the cranial and medial regions
et al., 2017). It should be emphasized that the occurrence of fibro- of the breast compared to those in normal chicken breasts (Baldi
sis in WB may not necessarily affect the texture properties of the et al., 2018; Kuttappan et al., 2017; Mudalal et al., 2014, 2015;
breast (Sihvo et al., 2017), and therefore, the detection by palpa- Zambonelli et al., 2016). However, in relation to the height in the
tion may be an intricate task. WB myopathy is generally detected caudal region, the WS and WS/SM presented a similar dimension
easier when the birds reach the age of slaughter (around 40 days) to the normal group, while the SM, WB, and WS/WB breasts
in the producing farms, or after slaughter in the slaughterhouses had a larger dimension (Baldi et al., 2018; Kuttappan et al., 2017;
(Papah et al., 2017). Mudalal et al., 2014; Mudalal et al., 2015; Zambonelli et al.,
The condition SM is characterized by a separation of the bundles 2016). No relationship was observed between breast width and
of muscle fibers mainly in the cranial region of the breast muscle myopathies (Baldi et al., 2018; Mudalal et al., 2014, 2015; Zam-
and this may, or may not, be associated with WS (Baldi et al., 2018). bonelli et al., 2016), yet, a largest length in breasts was observed
It is still to be determined whether this histological condition in WS, SM, and WS/SM affected muscles in comparison with
occurs in the living animal. normal breasts (Baldi et al., 2018; Mudalal et al., 2014). In WB,
The weakness in the technique for detecting these myopathies an accretion of extracellular material has been reported to occur in
is a key point and fast and nondestructive methods need to be lesion areas. According to Soglia et al. (2017) and Daalgard et al.
developed and applied. On this regard, some successful attempts (2018) this accumulation, which may concur with water mobility,
have been made, such as the application of radiofrequency spectra is significantly higher in breasts affected by WB than in normal
in the detection of WS in skin-on chicken carcasses (Traffano- ones. Therefore, the high incidence of myopathies, and particu-
Schiffo, Castro-Giraldez, Colom, & Fito, 2017) and the usage of larly WB, in large breast muscles, may be a consequence of the
near-infrared spectroscopy (NIR) for the detection of breast fillets pathological mechanisms involving edema and inflammation.
chicken with WB (Wold, Måge, Løvland, Sanden, & Ofstad, 2018; In addition to changes in size and weight of the pectoralis major
Wold, Veiseth-Kent, Høst, & Løvland, 2017). Although the use of muscles, degenerative lesions are also observed in WS, WB, and
NIR in the detection of WB was effective, the technique was de- SM affected cases. In detail, WS exhibits an increased deposition
veloped for breast fillet, and its efficacy in carcasses was not tested. of fat and proliferation of connective tissue within the endo- and
It is therefore not known to which extent skin would interfere peri-mysial compartments, loss of cross-striation, degeneration up
in the detection of the myopathy, limiting the application of this to necrosis and regeneration of the fibers, interstitial inflamma-
technique on skin-on carcasses. Other accurate methods, such as tion, edema, infiltration of inflammatory cells—macrophages and
histological or biochemical tests may require removing the animals lymphocytes; with the severity of the histopathological lesions
from the slaughter line and sampling the muscles which means the gradually increasing with the severity of the striae (Kuttappan
destruction of the carcass. The use of biomarkers to identify my- et al., 2013a,b). Sihvo et al. (2017) pointed out that broiler chick-
opathies in live birds by “omics” platforms using biological samples ens affected by WB myopathies begin to develop morphological
(that is, plasma) may be a highly remarkable option (further in- changes at approximately two weeks of age and, as the age in-
formation in Section 7; Final Remarks). In addition, there is also creases, the lesions tend to aggravate. The most frequent changes
a need to create a standard classification scale to identify progress observed in chicken breasts with WB are multifocal degenera-
in the degree of severity of WS, WB, and SM myopathies, which tion and necrosis, loss of striation, infiltration of macrophages and
may improve current classification methods. Some recent attempts lymphocytes in the fibers, thickening of the perimysial connective
have been made on this regard (Daalgard et al., 2018) tissue, fibrous connective tissue in superficial and deep part, de-
posit of extracellular collagen, myodegeneration, and regeneration
Morphometric and histological changes in abnormal (Sihvo et al., 2014; 2017), nuclear internalization, hyaline and vac-
chicken breasts uolar degeneration (Soglia et al., 2017), variability in shape, size,
Chickens with WS, WB, and SM usually present changes in and diameter of the muscle fibers, edema, vasculitis, and perivas-
morphometric measurements of the pectoralis major muscle (as cular infiltrations in veins (Sihvo et al., 2014, 2017). Overall, SM
briefly shown in Table 1), especially in breast weight, yield, and affected muscles display histological features similar to those pre-
height. Morphometric analyses have shown that all data concern- viously reported in WS and WB. However, the separation of the
ing breast dimensions (Alnahhas et al., 2016; Baldi et al., 2018; fiber bundles composing the muscle tissue as well as the progres-
Bowker & Zhuang, 2016; Brambila, Bowker, & Zhuang, 2016; sive degradation of the perimysial connective tissue observed by
Mudalal, Babini, Cavani, & Petracci, 2014; Mudalal, Lorenzi, Baldi et al. (2018) can be considered distinctive microscopic fea-
Soglia, Cavani, & Petracci, 2015), with the only exception of tures of SM defect. At last, it is worth mentioning that WS/WB
breast width (Baldi et al., 2018; Mudalal et al., 2014, 2015), were and WS/SM conditions generally displayed pathological charac-
affected by WS. Data indicate that these parameters increase with teristics similar to those observed in muscles affected by just one
the severity of striping thus confirming that the occurrence of WS myopathy.
is linked to thicker or heavier fillets. Similar results were reported
by Dalle Zotte et al. (2017), Mudalal et al. (2015), Xing et al. Physical–chemical modifications in abnormal chicken
(2017a), Xing, Zhao, Cai, Zhou, and Xu (2017b), and Dalgaard breasts
et al. (2018) in chicken meat affected by WB. Chatterjee, Zhuang, Chicken breast affected by either one or two of the myopathies
Bowker, Rincon, and Sanchez-Brambila (2016), Zambonelli et al. (WB, WS, and SM) display differences in chemical composition


C 2019 Institute of Food Technologists® Vol. 00, 2019 r Comprehensive Reviews in Food Science and Food Safety 5
Emerging broiler meat abnormalities . . .

Table 1–Variation in morphometric parameters of chicken breast affected by white striping (WS), wooden breast (WB), and spaghetti meat (SM)
myopathies.

Morphometric parameters Abnormality Occurrence References

Live weight WS moderate WS>N Alnahhas et al. (2016)
No effect Kuttappan et al. (2012b)
WS severe WS>N Alnahhas et al. (2016); Kuttappan et al. (2013b)
No effect Kuttappan et al. (2012b)
Breast weight WSc WS>N Baldi et al. (2018); Mudalal et al. (2015)
WS moderate WS>N Bowker and Zhuang (2016); Brambila et al. (2016)
No effect Kuttappan et al. (2012b)
WS severe WS>N Bowker and Zhuang (2016); Brambila et al. (2016);
Kuttappan et al. (2013b); Kuttappan et al. (2013b,
2012b); Mudalal et al. (2014)
WBd WB>N Dalle Zotte et al. (2017)
WB moderate WB>N Dalgaard et al. (2018)
WB severe WB>N Dalgaard et al. (2018); Xing et al. (2017b); Mudalal et al.
(2015)
SM SM>N Baldi et al. (2018)
WS/WB WS/WB>N Chatterjee et al. (2016); Zambonelli et al. (2016); Mudalal
et al. (2015)
WS/SM WS/SM>N Baldi et al. (2018)
Breast yield WS moderate WS>N Alnahhas et al. (2016); Kuttappan et al. (2012b)
WS severe WS>N Alnahhas et al. (2016); Kuttappan et al. (2012b); Kuttappan
et al. (2012b, 2013b, 2017ba )
No effect Kuttappan et al. (2017)b
WB severe WB>N Kuttappan et al. (2017)a,b
WS/WB WS/WB>N Kuttappan et al. (2017)a,b
Breast Length WSc WS>N Baldi et al. (2018)
No effect Mudalal et al. (2015)
WS severe WS>N Mudalal et al. (2014)
WB severe No effect Mudalal et al. (2015)
SM No effect Baldi et al. (2018)
WS/WB No effect Mudalal et al. (2015); Zambonelli et al. (2016)
WS/SM WS/SM>N Baldi et al. (2018)
Breast width WSc No effect Baldi et al. (2018); Mudalal et al. (2015)
WS severe No effect Mudalal et al. (2014)
WB severe No effect Mudalal et al. (2015)
SM No effect Baldi et al. (2018)
WS/WB WS/WB>N Zambonelli et al. (2016)
No effect Mudalal et al. (2015)
WS/SM No effect Baldi et al. (2018)
Breast top height WSc WS>N Baldi et al. (2018); Mudalal et al. (2015)
WS severe WS>N Mudalal et al. (2014), Kuttappan et al. (2017)a
No effect Kuttappan et al. (2017)b
WB severe WB>N Kuttappan et al. (2017)a ; Mudalal et al. (2015)
No effect Kuttappan et al. (2017)b
SM SM>N Baldi et al. (2018)
WS/WB WS/WB>N Kuttappan et al. (2017)a,b ; Mudalal et al. (2015);
Zambonelli et al. (2016)
WS/SM WS/SM>N Baldi et al. (2018)
Breast middle height WSc WS>N Baldi et al. (2018); Mudalal et al. (2015)
WS severe WS>N Mudalal et al. (2014)
WB severe WB>N Mudalal et al. (2015)
SM SM>N Baldi et al. (2018)
WS/WB WS/WB>N Mudalal et al. (2015); Zambonelli et al. (2016)
WS/SM WS/SM>N Baldi et al. (2018)
Breast bottom height WSc No effect Baldi et al. (2018); Mudalal et al. (2015)
WS severe No effect Mudalal et al. (2014); Kuttappan et al. (2017)b
WS>N Kuttappan et al. (2017)a
WB severe WB>N Kuttappan et al. (2017)a,b ; Mudalal et al. (2015)
SM SM>N Baldi et al. (2018)
WS/WB WS/WB>N Kuttappan et al. (2017)a,b ; Mudalal et al. (2015);
Zambonelli et al. (2016)
WS/SM No effect Baldi et al. (2018)
a Male Broiler 6 weeks-day.
b Male Broiler 9 weeks-day.
c Medium-to-thick white striations.
d Myopathy degree nonspecified.

when compared with normal chicken breasts (Table 2). WS ex- The content of moisture and ashes in affected breasts vary be-
hibits higher fat and collagen contents and lower protein level tween authors and studies (Baldi et al., 2018; Kuttappan et al.,
compared to normal meat (Baldi et al., 2018; Kuttappan, Brewer, 2012b; Petracci et al., 2014; Soglia et al., 2016a,b; Soglia et al.,
Apple, Waldroup, & Owens, 2012b; Malila et al., 2018; Mudalal 2018b). It is important to emphasize that although some studies
et al., 2014; Petracci, Mudalal, Babini, & Cavani, 2014; Soglia show an increase in collagen content in breasts with WS, this in-
et al., 2016a; Soglia, Laghi, Canonico, Cavani, & Petracci 2016b). crease is not observed in all reports. According to the literature, in

6 Comprehensive Reviews in Food Science and Food Safety r Vol. 00, 2019 
C 2019 Institute of Food Technologists®
Emerging broiler meat abnormalities . . .

Table 2–Variation in chemical parameters of chicken breast affected by white striping (WS), wooden breast (WB), and spaghetti meat (SM) myopathies.

Chemical parameters Abnormality Occurrence References

Proximate composition
Moisture WSd No effect Baldi et al. (2018)a ; Soglia et al. (2016a,b)
WS moderate No effect Soglia et al. (2018b)c ; Kuttappan et al. (2012b); Petracci
et al. (2014)
WS severe No effect Soglia et al. (2018b)c ; Kuttappan et al.(2012b); Petracci
et al. (2014)
WS>N Mudalal et al. (2014)
WB moderate WB>N Wold et al. (2017)
WB severe WB>N Wold et al. (2017); Cai et al. (2018); Soglia et al.
(2016a,b)
SM SM>N Baldi et al. (2018)a
WS/WB WB>N Soglia et al. (2016a,b)
WS/SM WS/SM>N Baldi et al. (2018)b
Protein WSd WS<N Baldi et al. (2018)a ; Soglia et al. (2016b)
WS moderate No effect Soglia et al. (2018b)c Kuttappan et al. (2012b)
WS<N Petracci et al. (2014)
WS severe No effect Soglia et al. (2018b)c
WS<N Petracci et al. (2014); Mudalal et al. (2014); Kuttappan
et al. (2012b)
WB moderate WB<N Wold et al. (2017)
WB severe WB<N Wold et al. (2017); Cai et al. (2018); Soglia et al.
(2016a,b)
SM SM<N Baldi et al. (2018)a
WS/WB WS/WB<N Soglia et al. (2016a,b)
WS/SM WS/SM<N Baldi et al. (2018)a
Lipid WSd WS>N Baldi et al. (2018)a ; Soglia et al. (2016b)
WS moderate No effect Soglia et al. (2018b)c ; Kuttappan et al. (2012b)
WS severe WS>N Soglia et al. (2018b)c ; Kuttappan et al. (2012b)
WB moderate No effect Wold et al. (2017)
WB severe WB>N Wold et al. (2017); Cai et al. (2018); Soglia et al.
(2016a,b)
SM No effect Baldi et al. (2018)a
WS/WB WS/WB>N Soglia et al. (2016a,b)
WS/SM WS/SM>N Baldi et al. (2018)b
Ash WSd No effect Baldi et al. (2018)a
WS moderate No effect Kuttappan et al. (2012b)
WS<N Soglia et al. (2018b)c
WS severe No effect Kuttappan et al. (2012b)
WS<N Soglia et al. (2018b)c ; Mudalal et al. (2014)
WB severe WS<N Soglia et al. (2016a)
SM SM<N Baldi et al. (2018)b
WS/WB WS<N Soglia et al. (2016a)
WS/SM WS/SM<N Baldi et al. (2018)b
Collagen WSd No effect Baldi et al. (2018)a ; Soglia et al. (2016b)
WS moderate WS>N Petracci et al. (2014)
No effect Soglia et al. (2018b)c
WS severe No effect Soglia et al. (2018b)c
WS>N Petracci et al. (2014); Mudalal et al. (2014)
WB severe No effect Cai et al. (2018)
WB>N Soglia et al. (2016a,b)
SM No effect Baldi et al. (2018)a
WS/WB No effect Soglia et al. (2016b)
WB>N Soglia et al. (2016a)
WS/SM No effect Baldi et al. (2018)a
Intramuscular fat WS moderate WS>N Petracci et al. (2014)
WS severe WS>N Petracci et al. (2014); Mudalal et al. (2014)
TBARS WSd No effect Soglia et al. (2016b)
WS moderate No effect Alnahhas et al. (2016)
WS severe No effect Alnahhas et al. (2016)
WB severe WB>N Soglia et al. (2016b)
WS/WB No effect Soglia et al. (2016b)
SFA WSd WS<N Soglia et al. (2016b)
WS severe WS<N Kuttappan et al. (2012b, 2013b)
WB severe No effect Soglia et al. (2016b)
WS/WB No effect Soglia et al. (2016b)
MUFA WSd No effect Soglia et al. (2016b)
WS severe WS>N Kuttappan et al. (2012b, 2013b)
WB severe No effect Soglia et al. (2016b)
WS/WB No effect Soglia et al. (2016b)
PUFA WSd No effect Soglia et al. (2016b)
WS severe No effect Kuttappan et al. (2012b)
WS<N Kuttappan et al. (2013b)
WB severe No effect Soglia et al. (2016b)
WS/WB No effect Soglia et al. (2016b)
EPA WSd No effect Soglia et al. (2016b)
(Continued)


C 2019 Institute of Food Technologists® Vol. 00, 2019 r Comprehensive Reviews in Food Science and Food Safety 7
Emerging broiler meat abnormalities . . .

Table 2–Continued.

Chemical parameters Abnormality Occurrence References

WS severe No effect Kuttappan et al. (2013b)
WS<N Kuttappan et al. (2012b)
WB severe No effect Soglia et al. (2016b)
WS/WB WS/WB<N Soglia et al. (2016b)
DPA WSd No effect Soglia et al. (2016b)
WS severe WS<N Kuttappan et al. (2012b, 2013b)
WB severe No effect Soglia et al. (2016b)
WS/WB WS/WB<N Soglia et al. (2016b)
DHA WSd No effect Soglia et al. (2016b)
WS severe WS<N Kuttappan et al. (2012b, 2013b)
WB severe No effect Soglia et al. (2016b)
WS/WB No effect Soglia et al. (2016b)
Carbonyls WSd No effect Soglia et al. (2016b)
WB severe WB>N Soglia et al. (2016b)
WS/WB WS/WB>N Soglia et al. (2016b)
a Superficial and deep position of breast meat.
b Superficial position of breast meat.
c Turkey breast meat.
d Medium-to-thick white striations.

approximately 50% of the articles published so far, this increase Impact of WB, WS, and SM on Meat Quality
was evidenced with the appearance and intensity of WS myopa- Depending on the severity of the defects, breast affected by the
thy. As examples, Baldi et al. (2018, 2019) and Soglia et al. (2016b) aforementioned myopathies may be withdrawn from food chain.
reported no effect of the WS condition on collagen content. Dif- A variable share of the breasts accepted as suitable for human con-
ferences were neither detected for turkeys with different degree of sumption will also account for economic losses owing to (i) con-
WS in regard to protein content and collagen (Soglia et al., 2018b). demnation/trimming (whole breast, carcass); (ii) decreased yield
Kuttappan et al. (2012b) and Petracci et al. (2014) pointed out that and value (that is, water-holding capacity [WHC], emulsifying
the protein content decreases as the degree of WS increases from and gelling abilities, and so on); (iii) increased need of manual
normal to severe and an opposite trend is observed for lipid con- sorting at deboning line (adding and training of personnel for
tent. Ash content was found to be lower in breast muscles with grading/sorting); and (iv) the rejection by consumers due to un-
moderate (Soglia et al., 2018b) and severe WS (Mudalal et al., desirable sensory properties. Despite of a likely yield reduction
2014; Soglia et al., 2018b). Alnahhas et al. (2016) and Soglia et al. due to increased drip and cook losses, processing altered breasts
(2016b) detected the absence of the effect of moderate and se- could be an alternative option to market altered breasts as fresh
vere WS meat on the TBARS content. Although lipid content meat as hence, avoid a potential rejection by consumer. Kuttappan
is generally higher in WS, these breasts do not seem to be more et al. (2016) roughly estimated in $200 million per year the cost of
susceptible to lipid oxidation than normal ones. Moreover, higher such conditions only in the United States. Given the remarkable
levels of monounsaturated fatty acids and lower levels of saturated, economic loss entailed by these emerging myopathies, the charac-
EPA, DPA, and DHA fatty acids were observed in chicken breasts terization of their quality profile and the underlying mechanisms
with severe WS compared to normal breasts (Kuttappan et al., of such impaired traits is essential to adequately ascribe breast mus-
2012b; Kuttappan et al., 2013a,b). cles to a suitable processing technology and minimize the negative
Similar findings have been observed in chicken breast with WB impact of the myopathy on chicken quality and consumer ac-
myopathy (with or without WS). Several authors have reported ceptability. The following lines will be devoted to describing the
consistent results including significant reduction in protein and ash quality of WB, WS, and SM samples in terms of functionality,
contents and increase in moisture and lipid levels in WB muscles sensory properties, and nutritional value. Some of these results are
compared to their normal counterpart (Cai et al., 2018; Soglia summarized in Table 3.
et al., 2016a,b; Wold et al., 2017). The collagen content in the
WB, SM, WB, WS/WB, and WS/SM presented contrasting re- Functionality
sults since some studies showed a higher content than the normal The functional properties of meat are typically ascribed to the
group and others did not observe a significant difference (Baldi ability of the myofibrillar proteins to hold water, emulsify lipids and
et al., 2018; Cai et al., 2018; Soglia et al., 2016a,b). Soglia et al. form stable gels. In turn, meat proteins functionality is dependent
(2016b) emphasized that even WB and WS/WB breasts showed on their amino acid composition, their three-dimensional struc-
no difference in saturated, monounsaturated, polyunsaturated, and ture and the complex fibrillar architecture in intact muscle (Pearce,
DHA fatty acids if compared to normal breasts; the former were Rosenvold, Andersen, & Hopkins, 2011). Biochemical processes
more susceptible to oxidation, with a higher level of carbonyls in affecting protein composition and integrity (proteolysis, oxidation,
WB and WS/WB breasts and a higher TBARS content in WS glycation, and so on) may impair their ability to interact with other
and WB breasts. They also reported that only breast WS/WB pre- biomolecules (lipids) and water (Estévez, 2011). The first reports
sented lower DPA and EPA contents. Changes in the chemical on the pathological conditions of WS and WB made by Kuttappan
composition of WB and WS/WB meats are consistent with the et al. (2012b) and Sihvo et al. (2014) were readily followed by the
replacement of fibers by adipose tissue (lipidosis) and the accre- description of their impaired quality traits and particularly, of their
tion of extracellular water as a result of edema and inflammatory poor ability to hold water (Petracci, Mudalal, Bonfiglio, & Cavani,
processes (Clark & Velleman, 2017; Kawasaki, Iwasaki, Yamada, 2013; Trocino et al., 2015). There is a consensus on the impaired
Yoshida, & Watanabe, 2018). functionality of WB and WS and that such adverse condition is

8 Comprehensive Reviews in Food Science and Food Safety r Vol. 00, 2019 
C 2019 Institute of Food Technologists®
Emerging broiler meat abnormalities . . .

Table 3–Technological properties of chicken breast with white striping (WS), wooden breast (WB), and spaghetti meat (SM).

Technological properties Abnormality Occurrence References

Lh WSh No effect Baldi et al. (2018); Mudalal et al. (2015); Tasoniero et al.
(2016); Trocino et al. (2015)
WS moderate WS>N Alnahhas et al. (2016)b
No effect Alnahhas et al. (2016)b ; Bower and Zhuang (2016);
Brambila et al. (2016); Petracci et al. (2013); Soglia
et al. (2018b)c ; Kuttappan et al. (2013c)
WS severe WS>N Alnahhas et al. (2016)b
No effect Alnahhas et al. (2016)b ; Bower and Zhuang (2016);
Brambila et al. (2016); Petracci et al. (2013); Soglia
et al. (2018b)c ; Kuttappan et al. (2013c)
WBi WB>N Dalle Zotte et al. (2017)
No effect Chen et al. (2018); Trocino et al. (2015)
WB moderate No effect Wold et al. (2017)
WB severe WB>N Cai et al. (2018); Wold et al. (2017)
No effect Mudalal et al. (2015); Xing et al. (2017b)
SM No effect Baldi et al. (2018)
WS/WB WS/WB<N Zambonelli et al. (2016)
No effect Chatterjee et al. (2016); Mudalal et al. (2015);
Tasoniero et al. (2016)
WS/SM No effect Baldi et al. (2018)
ah WSh No effect Baldi et al. (2018); Mudalal et al. (2015); Tasoniero et al.
(2016)
WS>N Trocino et al. (2015)
WS moderate WS>N Petracci et al. (2013)
No effect Alnahhas et al. (2016)a,b ; Bower and Zhuang (2016);
Brambila et al. (2016); Soglia et al. (2018b)c ;
Kuttappan et al. (2013c)
WS severe WS>N Petracci et al. (2013)
No effect Alnahhas et al. (2016)a,b ; Bower and Zhuang (2016);
Brambila et al. (2016); Soglia et al. (2018b)c ;
Kuttappan et al. (2013c)
WBi WB>N Dalle Zotte et al. (2017)
No effect Trocino et al. (2015)
WB moderate No effect Wold et al. (2017)
WB severe WB>N Cai et al. (2018); Wold et al. (2017); Xing et al. (2017b)
No effect Mudalal et al. (2015)
SM No effect Baldi et al. (2018)
WS/WB WS/WB>N Chatterjee et al. (2016); Tasoniero et al. (2016)
No effect Mudalal et al. (2015); Zambonelli et al. (2016)
WS/SM No effect Baldi et al. (2018)
bh WSh No effect Baldi et al. (2018); Mudalal et al. (2015); Tasoniero et al.
(2016)
WS<N Trocino et al. (2015)
WS moderate No effect Petracci et al. (2013); Alnahhas et al. (2016)a,b ; Bower
and Zhuang (2016); Brambila et al. (2016); Soglia
et al. (2018b)c ; Kuttappan et al. (2013c)
WS severe WS>N Petracci et al. (2013); Kuttappan et al. (2017)d,e
No effect Alnahhas et al. (2016)a,b ; Bower and Zhuang (2016);
Brambila et al. (2016); Soglia et al. (2018b)c ;
Kuttappan et al. (2013c)
WBi WB>N Dalle Zotte et al. (2017)
No effect Trocino et al. (2015)
WB moderate WB>N Wold et al. (2017)
WB severe WB>N Cai et al. (2018); Kuttappan et al. (2017)d,e ; Mudalal
et al. (2015); Wold et al. (2017)
No effect Xing et al. (2017b)
SM No effect Baldi et al. (2018)
WS/WB WS/WB>N Kuttappan et al. (2017)d,e ; Tasoniero et al. (2016)
No effect Chatterjee et al. (2016); Mudalal et al. (2015);
Zambonelli et al. (2016)
WS/SM No effect Baldi et al. (2018)
pH WSh WS>N Mudalal et al. (2015); Trocino et al. (2015)
No effect Tasoniero et al. (2016)
WS moderate WS>N Alnahhas et al. (2016)b ; Brambila et al. (2016); Bower
and Zhuang (2016)
No effect Petracci et al. (2013); Alnahhas et al. (2016)a ; Soglia
et al. (2018b)c ; Kuttappan et al. (2013c)
WS severe WS>N Petracci et al. (2013); Bower and Zhuang (2016)
No effect Alnahhas et al. (2016)a,b ; Brambila et al. (2016); Soglia
et al. (2018b)c ; Kuttappan et al. (2013c, 2017)d,e
WBi WB>N Dalle Zotte et al. (2017)
No effect Chen et al. (2018); Trocino et al. (2015)
WB moderate No effect Dalgaard et al. (2018)f ; Wold et al. (2017)
(Continued)


C 2019 Institute of Food Technologists® Vol. 00, 2019 r Comprehensive Reviews in Food Science and Food Safety 9
Emerging broiler meat abnormalities . . .

Table 3–Continued.

Technological properties Abnormality Occurrence References

WB severe WB>N Cai et al. (2018); Dalgaard et al. (2018)g ; Kuttappan
et al. (2017)d,e ; Xing et al. (2017b)
No effect Mudalal et al. (2015); Wold et al. (2017); Soglia et al.
(2016a)
WS/WB WS/WB>N Chatterjee et al. (2016); Kuttappan et al. (2017)d,e ;
Mudalal et al. (2015); Zambonelli et al. (2016); Soglia
et al. (2016a)
No effect Tasoniero et al. (2016)
Shear force WSh No effect Tasoniero et al. (2016); Trocino et al. (2015)
WS moderate No effect Alnahhas et al. (2016)a,b ; Soglia et al. (2018b)c ;
Kuttappan et al. (2013c); Petracci et al. (2013)
WS severe WS<N Petracci et al. (2013)
No effect Soglia et al. (2018b)c ; Alnahhas et al. (2018b)a,b ;
Kuttappan et al. (2013c)
WBi No effect Dalle Zotte et al. (2017)
WB>N Trocino et al. (2015)
WB moderate No effect Dalgaard et al. (2018)
WB severe No effect Cai et al. (2018); Dalgaard et al. (2018)
WS/WB WS/WB>N Chatterjee et al. (2016); Tasoniero et al. (2016);
Zambonelli et al. (2016)
Hardness WBi WB<N Chen et al. (2018)
WB moderate No effect Dalgaard et al. (2018)f,g
WB severe WB>N Dalgaard et al. (2018)f,g ; Soglia et al. (2016a)
WS/WB WS/WB>N Chatterjee et al. (2016); Soglia et al. (2016a)
Cook loss WSh WS>N Mudalal et al. (2015)
No effect Tasoniero et al. (2016); Trocino et al. (2015)
WS moderate No effect Alnahhas et al. (2016)a,b ; Bower and Zhuang (2016);
Soglia et al. (2018b)c ; Brambila et al. (2016);
Kuttappan et al. (2013c)
WS>N Petracci et al. (2013)
WS severe No effect Alnahhas et al. (2016)b ; Bower and Zhuang (2016);
Soglia et al. (2018b)c ; Brambila et al. (2016);
Kuttappan et al. (2013c)
WS>N Alnahhas et al. (2016)a ; Mudalal et al. (2014); Petracci
et al. (2013)
WBi WB>N Dalle Zotte et al. (2017); Trocino et al. (2015)
WB moderate WB>N Dalgaard et al. (2018)f,g
WB severe WB>N Dalgaard et al. (2018)f,g ; Xing et al. (2017b); Mudalal
et al. (2015); Soglia et al. (2016a)
WS/WB WS/WB>N Zambonelli et al. (2016); Mudalal et al. (2015); Soglia
et al. (2016a)
No effect Tasoniero et al. (2016)
Thaw loss WSh No effect Trocino et al. (2015)
WS moderate WS<N Bowker and Zhuang (2016)
WS severe WS<N Bowker and Zhuang (2016)
WBi No effect Dalle Zotte et al. (2017); Trocino et al. (2015)
Salt-induced water uptake WS moderate No effect Bowker and Zhuang (2016)
WS severe No effect Bowker and Zhuang (2016)
Drip loss WSi WS<N Mudalal et al. (2015)
WS moderate No effect Alnahhas et al. (2016)a,b ; Petracci et al. (2013)
WS<N Soglia et al. (2018b)c
WS severe WS>N Alnahhas et al. (2016)a
No effect Alnahhas et al. (2016)b ; Soglia et al. (2018b)c ;
Kuttappan et al. (2017)d,e ; Petracci et al. (2013)
WB moderate No effect Dalgaard et al. (2018)f,g
WB severe WB>N Cai et al. (2018); Dalgaard et al. (2018)g ; Xing et al.
(2017b); Kuttappan et al. (2017)d,e ; Mudalal et al.
(2015); Soglia et al. (2016a)
No effect Dalgaard et al. (2018)f
WS/WB WS/WB>N Kuttappan et al. (2017)d,e
No effect Zambonelli et al. (2016); Mudalal et al. (2015); Soglia
et al. (2016a)
a Broiler line with high ultimate pH.
b Broiler line with low ultimate pH.
c Turkey breast meat.
d Male Broiler 6 weeks-day.
e Male Broiler 9 weeks-day.
f Sample determined on the day 0.
g Sample determined on the day 1.
h Medium-to-thick white striations.
i Myopathy degree non-specified.

10 Comprehensive Reviews in Food Science and Food Safety r Vol. 00, 2019 
C 2019 Institute of Food Technologists®
Emerging broiler meat abnormalities . . .

more intense at higher severity degrees of the myopathy (Tijare tion. Since the oxidative damage to meat proteins has been found
et al., 2016). Among the three muscle lesion types, WB displays to impair their ability to hold water and form emulsions and gels
particularly poor functionality which is manifested in low yields, (Estévez, 2015; Utrera & Estévez, 2012), this condition may also
impaired marinade uptake and reduced ability to hold water un- contribute to impair the functionality of WB muscles. The im-
der both raw (increased drip loss) and heat treatment conditions pact of WS condition alone on meat functionality is subject of
(increased cooking loss) (Dalgaard et al., 2018; Dalle-Zotte et al., debate as contradictory results are found in literature. Kuttappan
2017; Kuttappan et al., 2017, Mudalal et al., 2015; Soglia et al., et al. (2013c) found no major effects of severe WS condition on
2016a). Bowker, Maxwell, Zhuang, and Adhikari (2018) recently protein functionality as measured by cooking loss. To similar con-
evaluated the performance of severe WB fillets during marination clusions came Bowker and Zhuang (2016) who observed similar
(0.75% NaCl/0.45 sodium phosphate) and oven-cooking (78 °C) myofibrillar protein solubility, water uptake, and cooking loss be-
and found that the marinade uptake and subsequent retention was tween WS and normal chicken breasts. Conversely, Mudalal et al.
lower in WB samples than in the normal counterparts. On the (2014) and Alnahhas et al. (2016), Petracci et al. (2013) found
same line, Aguirre, Owens, Miller, and Alvarado (2018) assessed higher cooking losses in severe WS breasts compared to normal.
the capacity of chicken muscles affected by severe WB condition Interestingly, the former authors aimed to explain the impaired
to retain marination solution injected in bulk with 15% brine WHC by reporting altered quantity (reduced concentration of
(0.55% NaCl in the brine, final concentration). Twenty minutes myofibrillar proteins) and quality (increased extent of carbonyla-
after injection, the authors observed a lower marinade retention tion) of proteins from WS muscles. Recently, Soglia et al. (2018b)
in severe WB samples than in normal ones (59% compared with characterized turkey breast muscles affected by WS condition at
83% of the marinade injected) and more intense loss of water different degrees of severity and, despite of the differences in size
(23% compared with 20%) upon a flat-top grill cooking proce- and chemical composition, other quality traits, including WHC,
dure (176 °C). Similar results were reported by Brambila, Bowker, remained unaffected by the myopathy. As hypothesized by the
Chatterjee, and Zhuang (2018) for severe WB muscles stored at authors, species-specific physiological responses to the genetic se-
either 4 °C or 20 °C prior to cooking at 76 °C. Regardless of lection in turkeys may explain the occurrence of WS condition
storage temperature, cooking losses of WB fillets were between in these birds with limited effects on meat quality. The limited
8% and 10% units higher than those of normal chicken breasts. knowledge available on the quality traits of SM was recently re-
Chen et al. (2018) prepared meat batters and meatballs from nor- ported by Baldi et al. (2018) who compared this myopathy with
mal, PSE-like and wooden chicken breasts. The poorer WHC in WS. A nuclear magnetic resonance relaxation analysis revealed a
PSE-like and WB samples occurred along with a larger proportion higher proportion of extra-myofibrillar water in the superficial
and mobility of free water as measured by NIR. Xing et al. (2017a) section of SM samples, which was manifested in a reduced WHC.
not only reported the impaired WHC of meat batters produced These samples also had higher concentration of oxidized proteins
from WB muscles; these authors observed inferior gelation prop- than normal breast muscles supporting a likely influence of the
erties in these batters compared to those produced from normal chemical state of meat proteins on their functionality.
breast muscle, which formed more regular and pored networks.
The same authors stated that increasing NaCl contents from 0% to Sensory properties
2% to 3% to 4% could improve the gelling abilities of WB batter Appearance, texture and flavor are the most appreciated quality
to even compare the gel properties of normal batters (Xing, Zhao, traits by consumers of chicken meat (Carvalho et al., 2017; Estévez,
Cai, Zhou, & Xu, 2017b). NaCl increases the ionic strength of the 2015). The muscle tissue degeneration undergone during the on-
environment and hence, facilitates the solubility and functionality set of the myopathies under study has a straightforward influence
of myofibrillar proteins (Lobo et al., 2016). Nevertheless, the well- on the relevant sensory traits, namely color and texture. The over-
established theories explaining the relationship between pH and all appearance of the breasts affected by the myopathies reported in
WHC in meat systems are not applicable to chicken breast muscles this review is clearly altered as compared to a normal breast muscle
affected by the myopathies under study. A low final pH (close to (Figure 2) and that is a direct consequence of the histological hall-
the isoelectric point of muscle proteins approximately 5.0) leads marks of each myopathy: presence of out-bulging and pale areas
to muscles with poor WHC while muscles with high final pH of hardened consistency in WB, appearance of white striations of
display better abilities to hold water (Hamm, 1961). However, the variable thickness parallel to the muscle fibers direction in WS,
ultimate pH of WB (Kuttappan et al., 2017) and breasts affected and the disintegration of the muscle tissue in fiber bundles in SM.
by WS (Petracci et al., 2013) and SM (Baldi et al., 2018, 2019) Instrumental measurement of the color of WB typically results
is typically several decimals (0.2 to 0.4) higher than normal breast in elevated lightness (L∗ ) (Dalle Zotte et al., 2017) and increased
muscles and yet, their WHC is lower. The severe degeneration yellowness (b∗ ) (Kuttappan et al., 2017) with this alterations be-
of the muscle tissue in these myopathies would explain the poor ing worsened at higher degrees of severity (Tasoniero, Cullere,
WHC and the lack of correspondence between this parameter Cecchinato, Puolanne, & Dalle Zotte, 2016). Discoloration may
and pH. On this line, Dalgaard et al. (2018) attributed to a dis- also be noticeable upon cooking as recently reported by Zhuang
rupted and abnormal tissue structure the divergence between pH and Bowker (2018) who found that the surface of cooked WB
and WHC in WB muscles. The accretion of interstitial matter fillets was darker, redder, and more yellow than that of fillets with-
in WB myopathy (water, collagen, and proteoglycans) may also out the WB condition. Also, recently, Baldi et al. (2018) observed
contribute to explain the water losses of these abnormal muscles that color modifications in WB (increased L∗ and b∗ values) and
upon compression and heating. At a molecular level, proteins have WS (increased b∗ and reduced redness, a∗ ) only occurred in the
also been found to be affected by the underlying oxidative stress superficial layer of the muscle (0.3 cm) while appearance of the
occurred during the onset of WB. Soglia et al. (2016b) found muscles was normal at deeper layers.
a timely connection between the extent of protein oxidation (as The impact of the myopathies on the textural properties of
measured by total protein carbonyls) in WB and WB/WS samples chicken breast muscles has been profusely documented by studies
and the proportion and mobility of extra-myofibrillar water frac- applying both instrumental and sensory assessments. Sun, Koltes,


C 2019 Institute of Food Technologists® Vol. 00, 2019 r Comprehensive Reviews in Food Science and Food Safety 11
Emerging broiler meat abnormalities . . .

Coon, Chen, and Owens (2018) found positive and significant moderate WS breasts. In agreement, trained panelists were able to
correlations between compression force and the severity of WB find differences only between severe WS samples and normal ones
condition (r = 0.79) in raw breast muscles, clearly incriminat- in the study carried out by Brambila et al. (2016), while moderate
ing the histological degeneration of muscle in its altered texture WS samples were identified as normal is terms of textural proper-
properties. Soglia et al. (2017) reported that the enlargement of ties. In particular, cooked (78 °C) breast fillets affected by severe
extracellular matrix and fibrosis might contribute in explaining WS received higher scores for cohesiveness, hardness, and chewi-
the different texture properties between superficial and deep lay- ness than normal and moderate WS samples. In the study of Baldi
ers in WB samples, with the superficial part exhibiting a higher et al. (2019), SM seemed to display a softer texture after cooking
amount of larger particles and a higher compression force com- which the authors attributed to a small content of collagen in these
pared to deeper layers. Aguirre et al. (2018) found correspondence samples compared to normal muscles and breast muscles affected
between texture profile analysis (TPA) and sensory evaluation of by WS and WB. Beyond the collagen content, several attempts
severe WB muscles irrespective of the cooking methods applied to understand the underlying causes for these texture alterations
(flat-top grill or oven, both at 176 °C). The altered muscles dis- have led scientists to establish connections between texture mea-
played higher scores for hardness, cohesiveness, denseness, chewi- surements and other chemical components (that is, water content
ness, springiness, crunchiness and fibrousness than normal chicken and mobility) and assorted biochemical processes (that is, prote-
breasts. Consistently, Brambila et al. (2018) found higher scores of olysis and oxidation). Soglia et al. (2018a) studied proteolysis and
cohesiveness and springiness in WB fillets cooked at 76 °C than calpain activity in WB muscles and concluded that the increased
normal counterparts. These authors also reported that springi- hardness in such myopathy seemed not to be exclusively attributed
ness, hardness, and fibrousness were perceived differently between to differences in the proteolytic processes taking place within the
ventral and dorsal sections of cooked WB fillets. Using two in- postmortem period. Tasoniero, Bertram, Young, Dalle Zotte, and
strumental texture techniques, namely, Meullenet-Owens Razor Puolanne (2017) investigated the relationship between hardness in
Shear (MORS) and TPA, Chatterjee et al. (2016) stated that fillets WB and the myowater properties in these altered breast muscles
affected by WB condition required more force to cut through and using nuclear magnetic resonance. While a connection between
were harder and chewier than normal chicken breast. These dif- increased muscle hardness and longer relaxation time of water
ferences between normal and WB breast muscles were consistent trapped into the myofibrillar matrix was found, the authors con-
regardless of the freshness (fresh compared with frozen-thawed) cluded that the role of myowater on muscle hardness was not fully
and the cooking (raw compared with cooked at 78 °C). Similar clarified by this study. In fact, changes in water compartmental-
results were reported by Tijare et al. (2016) using the MORS ization associated to proteolysis during tenderization process in
technique. Conversely, Soglia et al. (2017) stated that texture dif- pork and sheep meat (Straadt, Rasmussen, Andersen, & Bertram,
ferences between normal breasts and those affected by WB were 2007), seemed not to be applicable in chicken breast muscles.
mainly detected in raw meat, with the WB samples showing the Therefore, the underlying mechanisms behind the increased hard-
highest compression values. According to the authors, the ther- ness in WB muscles seemed to be an open topic that may be
mally labile nature of the cross-links will account for the com- covered by looking into the quantity and chemical state of meat
parable shear-force values with normal breast when measured on proteins. Given the proven oxidative stress occurred in the my-
cooked samples, despite of the increased amount of connective opathies and the close connection between protein oxidation and
tissue in the WB. When differences were detected in cooked sam- meat texture (Lund, Lametsch, Hviid, Jensen, & Skibsted, 2007),
ples, strategies have been proposed to alleviate the texture prob- this is a well-deserved aspect to be examined.
lems. Vacuum-tumbling marination lessened but not eliminated
the texture differences between normal and severe WB cooked Nutritional value
chicken, since the latter were harder, chewier and more fibrous The impact of WB, WS, and SM conditions on the nutritional
than the former (Maxwell, Bowker, Zhuang, Chatterjee, & Ad- value of the chicken meat has only been partially covered. Most
hikari, 2018). Furthermore, the impaired texture of WB muscles studies infer the nutritional value of the abnormal breast meats by
may also be noticeable after mincing and processing as exposed interpreting the changes in the chemical composition as compared
by Chen et al. (2018), who found lower instrumental hardness, to normal chicken breast muscle. As commented in the previous
cohesiveness and chewiness in cooked meatballs produced from section, the three myopathies have been found to display a higher
WB as compared to those produced from normal chicken meat. amount of fat and moisture and less protein than normal breast
A scanning electronic microscopy analysis of samples revealed that muscles (Baldi et al., 2019; Petracci et al., 2014). This variation
WB meatballs had networks with large aggregates and big cavities. in proximate composition leads to higher energy content in se-
While Chen et al. (2018) attributed these results to protein de- vere WS breast muscles than in normal ones (451 compared with
naturation processes and lack of functional myofibrillar proteins, 421 Kj/100 g) (Petracci et al., 2014). Slight modifications in the
the oxidation state of such proteins (not measured in that study) fatty acid compositions have been reported but no variations in
may not be ruled out given the close relationship between the major nutritional indexes suggest that lipid quality is similar be-
oxidative damage to proteins and meat protein aggregation and tween normal chicken breasts and those affected by the myopathies
impaired functionality (Estévez, 2011; Utrera & Estévez, 2012). under examination. Conversely, the protein quality is clearly wors-
The texture properties of WS breasts are not so affected as those ened in breast muscles affected by myopathies due to (i) the higher
with the WB condition (Baldi et al., 2018; Tasoniero et al., 2016). collagen content and lower of myofibrillar proteins (Petracci et al.,
In fact, several authors found negligible effects of the WS condi- 2014) and (ii) the increased concentration of oxidized proteins as
tion on the texture of chicken (Kuttappan et al., 2013c) and turkey measured by the total amount of protein carbonyls (Baldi et al.,
breasts (Soglia et al., 2018b). Other studies only reported signifi- 2019; Soglia et al., 2016b). Protein carbonylation is responsible
cant differences when the most severe degrees of WS were assessed. for the subsequent formation of insoluble aggregates and has also
Using the Allo-Kramer technique, Petracci et al. (2013) found been linked to an impaired in vitro digestibility of oxidized proteins
lower shear-force values in severe WS samples than in normal and (Estévez, 2015; Soladoye, Juárez, Aalhus, Shand, & Estévez, 2015).

12 Comprehensive Reviews in Food Science and Food Safety r Vol. 00, 2019 
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Emerging broiler meat abnormalities . . .

To our knowledge, the digestibility of proteins from WB, WS or meat received lower scores than normal chicken breasts due the
SM has never been assessed so far. Finally, no harmful chemical toughness feeling in the mouth.
species have been found in breast muscles affected by myopathies,
so no specific chemical hazard linked to these abnormal breast Means of Alleviation: Facts and Challenges
muscles can be envisaged. However, the three myopathies have Given the notable incidence of WB, WS, and SM and assorted
been associated with increased levels of lipid and protein oxida- negative consequences of such myopathies on chicken quality and
tion products with some of these species (that is, MDA or pro- consumers’ preferences, the search for solutions to avoid the oc-
tein carbonyls) being identified as potentially noxious to humans currence or alleviate the symptoms has become a primary objective
(Esterbauer, Muskiet, & Horrobin, 1993; Estévez & Luna, 2017). for animal and food scientists. To succeed in this venture, a previ-
ous identification of the pathological pathways of these myopathies
Consumer Perception is obviously required. Several studies have been conducted to as-
Consumers are the final recipients of chicken meat and chicken certain the possible factors involved in the occurrence of WS, WB,
products. The ultimate purpose of any livestock and agricultural and SM in fast-growing broilers (Table 4). It is widely accepted
production should be to fulfil consumer’s demand on poultry that the incidence of breast abnormalities rises with increasing
meat in terms of safety, sensory properties, nutritional value, and slaughter weight (Cruz et al., 2016; Lorenzi et al., 2014; Papah
animal welfare (Carvalho et al., 2017; Lusk, 2018; Magdelaine et al., 2017), growth rate (Kuttappan et al., 2012a, 2013a, 2017;
et al., 2008). However, the selection of fast-growth hybrids, the Lorenzi et al., 2014), and genetic potential for breast meat yield
optimization of conversion rates and high yields and subsequent (Alnahhas et al., 2016; Bailey et al., 2015; Livingston et al., 2018;
occurrence of the myopathies described in the present article, Lorenzi et al., 2014; Petracci et al., 2013; Trocino et al., 2015)
demonstrate that consumer’s expectation and animal welfare in agreement with strong genetic correlations found by Alnah-
may have not been in the focus of producers. The appearance has et al. (2016). This has been also confirmed by histological
of “altered meat” (that is, discoloration, petechiae, abnormal observations which evidenced that myodegeneration progress as-
marbling, disintegration of meat structure, bulges) may be enough sociated with the development of breast abnormalities is strictly
motivation for the consumer to reject breast meat affected by a related to age at slaughter (Griffin et al., 2018; Kawasaki et al.,
myopathy. However, the awareness of the underlying causes of 2018; Papah et al., 2017; Radaelli et al., 2017; Sihvo et al., 2017)
that altered breast meat and the identification of that product as and breast growth pattern (Papah et al., 2017) (Table 5; Radaelli
a “sick muscle” may consolidate the consumer rejection, even if et al., 2017). Recently, it was also demonstrated that egg storage
some of them would not particularly care about animal welfare. duration before hatching as well as manipulation of embryonic
While this reasonable hypothesis needs to be proven, it is obvious development by egg incubation temperatures and chick weight at
that internationally recognized media has recently been informing hatching can affect muscle morphology traits and related to occur-
of the occurrence of WS, their causes and consequences (The rence of breast abnormalities (Clark, Walter, & Velleman, 2017;
Sun, 2017). Even if the information divulged is (at times) biased Livingston et al., 2018). Lately, it was also hypothesized that there
and not fully scientifically legitimate, it is a fact that consumers is progression from WS to WB with increasing age at slaughter
from developed countries are increasing concerned on how (Griffin et al., 2018). Moreover, there are significant gender dif-
livestock are farmed and meat is produced (The Independent, ferences especially in WS and WB occurrence which appears to
2017). Consumer’s demanding attitude is reflected in that fact that be higher in males (Kuttappan et al., 2013a; Lorenzi et al., 2014;
chicken consumers in the United States are currently willing to Trocino et al., 2015). Otherwise, it was recently reported that SM
pay more for “slow growth chicken” meat as long as appropriate condition is more prevalent in female birds (Soglia, Mazzoni, &
information is provided in the label (Lusk, 2018). Petracci, 2019).
The most complete study of consumer’s attitude toward chicken While genetic selection and fast growth seem to be major causes
breast affected by myopathies was carried out by Kuttappan et al. for the occurrence of these myopathies, most of the studies aimed
(2012c) with U.S. consumers and several degrees of WS. In a hedo- to mitigate occurrence and severity of breast abnormalities have
nic assessment of overall linking for appearance, consumers’ scores been conducted in the field of animal nutrition (Table 4). The
significantly decreased as the severity of the myopathy increased main strategies involved reduction of dietary intake of energy or
(6.9, normal; 6.1, moderate WS; and 4.5 severe WS in a 9-point amino acids by either feed restriction or changing feed formula-
scale). Overall, almost 57% of the consumers clearly disliked fillets tion (Cruz et al., 2016; Kuttappan et al., 2013a; Livingston et al.,
with severe WS. The average purchase intent also significantly de- 2019; Livingston, Ferket, Brake, & Livingston, 2019; Meloche,
creased in breast affected by WS from 3.6 (normal) to around 2.5 Fancher, Emmerson, Bilgili, & Dozier, 2018a; Sachs et al., 2018;
(WS) in a 5-point scale. When requested to explain the reasons Trocino et al., 2015,b,c,d; Zampiga et al., 2018). However, over-
for their preferences, consumers highlighted the higher marbling all a reduction in the occurrence of breast muscle abnormalities
appearance in WS fillets, which may have been perceived as ab- was observed almost exclusively as an indirect result of decreased
normal in chicken breast. Dalle Zotte et al. (2017) also made a growth rate, slaughter weight, and/or breast yield (Cemin et al.,
worthy attempt to evaluate the impact of WB appearance on the 2018; Cruz et al., 2016; Livingston et al., 2018; Meloche, Dozier,
quality that consumers may perceive. According to this study, WB Brandebourg, & Starkey, 2018d; Sachs et al., 2018). Only a min-
myodegeneration worsens meat quality traits and the visual ap- imal dietary feed restriction (95%) and short-term reduction of
pearance of the affected breasts through the occurrence of bulges, lysine have allowed to slightly reduce severity of WS and WB, but
petechiae, and exudate. Given that the visual aspect of WS and adoption of these strategies under commercial condition are chal-
WB would affect product rejection at purchase (Kuttappan et al., lenging (Meloche et al., 2018d). Indeed, compensatory growth
2012c), Tasoniero et al. (2016) suggested whether a previous cook- following early feed restriction may even increase incidence of
ing treatment would avoid the negative influence of macroscopic breast abnormalities (Trocino et al., 2015). Analogously, differ-
lesions on consumer’s attitude. While this was efficiently accom- ent coccidiosis control approaches (Dalle Zotte, Tasoniero, Russo,
plished, a subsequent hedonic assessment revealed that WS/WB Longoni, & Cecchinato, 2015) and dietary supplementation with


C 2019 Institute of Food Technologists® Vol. 00, 2019 r Comprehensive Reviews in Food Science and Food Safety 13
Emerging broiler meat abnormalities . . .

Table 4–Factors affecting occurrence of white striping (WS), wooden breast (WB), and spaghetti meat (SM) in fast-growing broilers.

Factor Abnormality Occurrence References

Bird and live growth factors
Strain WS High > Standard breast-yield Petracci et al. (2013)
WS Lorenzi et al. (2014)
WS/WB Bailey et al. (2015)
WS/WB Trocino et al. (2015)
WS/WS Alnahhas et al. (2016)
Livingston et al. (2018)
Egg storage WS/WB Long > Short Livingston et al. (2018)
(only on WS in feed restricted
birds)
Gender WS Males > Females Kuttappan et al. (2013a)
WS Lorenzi et al. (2014)
WS/WB Trocino et al. (2015)
Body weight/age at WS High > Low Lorenzi et al. (2014)
slaughter WS/WB Cruz et al. (2016)
WB Papah et al. (2018)
Growth rate WS Fast > Slow Lorenzi et al. (2014)
WS/WB Kuttappan et al. (2012a, 2013a, 2017)
Dietary means
Phase-feeding WS No effect Kuttappan et al. (2013a)
Early dietary restriction WS/WB No effect Trocino et al. (2015)
Dietary restriction (95% WS/WB Freely > Feed restricted Meloche et al. (2018a)
freely)
Dietary restriction (90% WS/WB Freely > Feed restricteda Livingston et al. (2018)
freely)
Dietary energy and/or WS/WB No effect Meloche et al. (2018c)
amino acid density Bodle et al. (2018)
Lysine WS/WB High > Lowa Cruz et al. (2016)
Meloche et al. (2018d)
Lysine (short-term WS/WB Moderate effect on WS/WB Meloche et al. (2018b)
reduction) severitya
Arginine:lysine WS/WB No effect Bodle et al. (2018)
WS/WB/SM Low > High Zampiga et al. (2018)
Arginine WS/WB High > Lowa Livingston et al. (2018)
Glutamine WS/WB High > Lowa Livingston et al. (2019)
Methionine WS Synthetic > Natural sourcea Sachs et al. (2018)
Guanidinoacetic acid WS/WB/SM Slight effect on WB severity Cordoba-Noboa et al. (2018a,b)
Vitamin E WS No effect Kuttappan et al. (2012b)
Vitamin C WS/WB High > lowa Bodle et al. (2018)
Vitamin premix WS/WB No effect Bodle et al. (2018)
Selenium WB No effect Sihvo et al. (2017)
WS/WB Organic > Inorganica Cemin et al. (2018)
Organic trace minerals WS/WB/SM No effect Sirri et al. (2016)
Coccidiosis control WS No effect Dalle Zotte et al. (2015)
Postmortem factors
Chilling time WS No effect Kuttappan et al. (2013a)
a Changes result as indirect result of reduced growth rate, slaughter weight and/or breast yield.

Table 5–Factors affecting myodegeneration associated with occurrence of emerging abnormalities in fast-growing broilers.

Factor Degree of myodegeneration References

Egg incubation temperature High > Standarda Clark et al. (2017)
Hatching time Early > Latea Clark et al. (2017)
Gender No effect Radaelli et al. (2017)
Early dietary restriction Freely >Feed restricted Radaelli et al. (2017)
Body weight/age at slaughter Increased with age/body weight Radaelli et al. (2017), Sihvo et al.
(2017), Kawasaki et al. (2018),
Griffin et al. (2018), Papah et al.
(2018)
Breast yield/size Increased with pectoralis major Radaelli et al. (2017)
yield/size Papah et al. (2018)
a Changes result as indirect result of reduced growth rate, slaughter weight and/or breast yield.

antioxidants (vitamin E, C, and selenium) and organic trace min- that an increase of arginine:lysine ratio can have a mitigation effect
erals (Kuttappan et al., 2012b; Sihvo et al., 2017; Sirri et al., on breast meat abnormalities (Zampiga, Soglia, Petracci, Meluzzi,
2016), which overall have been supposed to reduce muscle fiber & Sirri, 2018) though a similar study led to a divergent outcome
oxidative stress associated with breast abnormality progress, did (Bodle et al., 2018). Therefore, until now it seems that there is
not result in any true mitigation effects. On the other hand, a lack of practical nutrition and management interventions to re-
slight reduction of moderate cases of WB has been obtained by duce growth-related abnormalities in the broiler industry without
dietary supplementation of guanidinoacetic acid used a precursor negatively affecting live and slaughter performances. Finally, few
of creatine, but strong interaction with diet composition was found studies were conducted to ascertain influence of slaughter phases
(Cordoba-Noboa et al., 2018a,b). Recently, it was demonstrated (Kuttappan et al., 2013a) which anyway appear to be of little

14 Comprehensive Reviews in Food Science and Food Safety r Vol. 00, 2019 
C 2019 Institute of Food Technologists®
Emerging broiler meat abnormalities . . .

Table 6–Processing solutions for alleviating meat quality consequences of occurrence of white striping (WS), wooden breast (WB), and spaghetti meat
(SM).

Meat processing
Abnormality solutions Effectiveness References
WB Breast fillet portioning high Bowker and Zhuang (2016)
WS/WB/SM (Separation of dorsal Baldi et al. (2018)
WB and ventral Bowker et al. (2018)
portions)
WS Vacuum tumbling poor Petracci et al. (2013)
WS/WB (Whole muscle) Mudalal et al. (2015)
WS/WB Soglia et al. (2016a))
WS/WB Tijare et al. (2016)
WB Bowker et al. (2018)
WB Maxwell et al. (2018)
WB Coarsely mincing high Sanchez-Brambila et al.
(Cooked patties) (2017, 2018)
WB Finely mincing High Xing et al. (2017b)
(Meat batters and Poor Chen et al. (2018)
meat balls)

importance in determining breast abnormalities (Petracci et al., and metabolomic profiles that may be employed for diagnosis pur-
2015). poses and ultimately, for a better comprehension of the mechanistic
Alternatively, some authors proposed meat processing solutions insight into these myopathies. Likewise, the application of genetic-
to minimize implications on product quality as summarized in based analysis is of interest given the clear genetic determinism of
Table 6. There are margins to mitigate undesirable effects on tech- some of these myopathies (Alnahhas et al., 2016). In this regard,
nological properties of processed products when abnormal meats Pampouille et al. (2018) recently carried out a genome-wide asso-
are included in the formulation of grounded and finely commin- ciation study in which a quantitative trait loci (QTL) mapping in
uted meat products (Brambila et al., 2018; Brambila, Chatterjee, WS revealed a polygenic inheritance of the defect and identified
Bowker, & Zhuang, 2017, Chen et al., 2018; Xing et al., 2017a), several candidate genes implicated in the muscular dystrophies. In
however significant quality reduction can arise when high-quality addition, Hubert, Williams, and Athrey (2018) by comparing birds
processed products (that is, enhanced whole-muscle and ground having fast- and slow-growth genetic backgrounds stated that WB
products) are manufactured by using especially WB abnormal raw is a potentially polygenic, complex syndrome, with molecular sim-
meat (Bowker, Maxwell, Zhuang, & Adhikari, 2018; Maxwell ilarities to neoplastic disorders. Vignale et al. (2017) contributed to
et al., 2018; Mudalal et al., 2015; Petracci et al., 2013; Soglia et al., understanding the impair protein and fat metabolism in chickens
2016a; Tijare et al., 2016). In addition, latest studies agree that affected by WS analyzing the expression of genes such as MuRF1,
overall manifestation of muscle abnormalities mainly affects the atrogin-1, IGF-1, insulin receptor (IR), fatty acid synthetase, and
superficial section of pectoralis major muscle, while the deep sec- acetyl CoA carboxylase (ACC).
tion was less affected (Baldi et al., 2018, 2019; Bowker & Zhuang, A fast progress is envisaged in the upcoming years and answers
2016; Bowker et al., 2018). Therefore, one possible strategy to to some unresolved questions may be found. Yet, some current
optimize its use after downgrading could be to separately process and urgent challenges are calling for attention such as the man-
the superficial (ventral) and deep (dorsal) layer of the abnormal agement of the high numbers of chicken breast muscles affected
pectoral muscle for exploiting their distinctive traits (Baldi et al., by WS, WB, and SM. One major issue is the early detection
2018, 2019; Bowker et al., 2018). and objective classification of the myopathies using nondestruc-
tive techniques. Aforementioned efforts have been made using
Final Remarks radiofrequency spectra (Traffano-Schiffo et al., 2017) and NIR to
The final words from this comprehensive review will be de- detect and grade WB and WS myopathies. Further advanced op-
voted to identifying unresolved problems and potential fields of tions are available as the fusion of Optical Coherence Tomography
study. The poultry sector is nowadays in continuous evolution in (OCT) and hyperspectral imaging proposed by Yoon, Bowker,
relation to the needs from the industry and the society. It seems and Zhuang (2017) or other imaging techniques already used
that the poultry industry as a whole (not only breeding compa- for an efficient non-destructive assessment of meat products such
nies) can no longer postpone long-anticipated decision-makings as ultrasounds (Corona, Garcı́a-Pérez, Santacatalina, Ventanas, &
and confront the awkward triangle formed by the selection of Benedito, 2014) and Magnetic Resonance Imaging (MRI) (Ca-
meat-type genotypes, fast muscle growth, and impaired quality ballero et al., 2018). Once the carcasses are identified as affected
in chicken breasts. Scientists, on the other hand, endure in their by the myopathies in a particular degree, decisions on the fate
commitment in searching for means of alleviation. Apparently, no of these chicken meats have to be taken. As consumers’ aware-
efficient solutions have been identified and hence, controlling the ness on the occurrence and origin of these chicken defects is
high incidence of these myopathies is a major commitment for increasing, transforming abnormal breasts into processed chicken
animal scientists. A profound understanding of the molecular basis products may be a feasible option. However, means to improve
of the underlying pathological processes is essential to establish their technological, sensory, and nutritional properties are also re-
well-reasoned and reliable strategies. On this line, the application quired. The increase in breast meat abnormalities is favoring the
of advanced methodologies based on omics platforms are of great transition from standard chicken to differentiated/price-premium
assistance. The study of the proteome in breasts affected by quality products (that is, animal welfare friendly, environment friendly,
defects (Schilling et al., 2017) and the metabolomic studies carried consumer health, buy local, and so on). In this unstable context,
out by Abasht et al. (2016) and Boerboom et al. (2018) in WB innovative strategies are also required to continuously adapt to this
and WS, respectively, contributed to establish unique proteomic mutable scenario and for this reason, know-how concerning the


C 2019 Institute of Food Technologists® Vol. 00, 2019 r Comprehensive Reviews in Food Science and Food Safety 15
Emerging broiler meat abnormalities . . .

whole production chain (from farm to fork) is more crucial than the woody breast condition. Poultry Science, 96(9), 3489–3494.
ever. https://doi.org/10.3382/ps/pex118
Brambila, G. S., Bowker, B. C., Chatterjee, D., & Zhuang, H. (2018).
Descriptive texture analyses of broiler breast fillets with the wooden breast
Acknowledgments condition stored at 4°C and –20°C. Poultry Science, 97, 1762–1767.
Marta Madruga, Leila Carvalho, and Elza Ida acknowledge the https://doi.org/10.3382/ps/pew327
CNPq- Conselho Nacional de Desenvolvimiento Cientifico e Carvalho, R. H., Ida, E. I., Madruga, M. S., Martı́nez, S. L., Shimokomaki,
Tecnológico (430832/2016-8). Mario Estévez acknowledge the M., & Estévez, M. (2017). Underlying connections between the redox
system imbalance, protein oxidation and impaired quality traits in pale, soft
CNPq and the Spanish Ministry of Economics and Competitive- and exudative (PSE) poultry meat. Food Chemistry, 215, 129–137.
ness (SMEC) (projects 474300/2011-0 and AGL2017-84586-R, https://doi.org/10.1016/j.foodchem.2016.07.182
respectively). Caballero, D., Antequera, T., Caro, A., Amigo, J. M., ErsbØll, B.K, Dahl, A.
B., & Pérez-Palacios, T. (2018). Analysis of MRI by fractals for prediction of
sensory attributes: A case study in loin. Journal of Food Engineering, 227,
Authors’ Contributions 1–10. https://doi.org/10.1016/j.jfoodeng.2018.02.005
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