Consumer Acceptability of Intramuscular Fat

Damian Frank1, Seon-Tea Joo2, Robyn Warner*
Author Information & Copyright
Faculty of Veterinary and Agricultural Science, University of Melbourne, Parkville, 3010, Australia
1CSIRO, Agriculture & Food, 11 Julius Ave North Ryde, New South Wales, 2113, Australia
2Division of Applied Life Science (BK21+), Gyeongsang National University, Jinju 52852, Korea
*Corresponding author: Robyn Warner, Faculty of Veterinary and Agricultural Science, University of Melbourne, Parkville, 3010, Australia. Tel: +61-3-9035-6663, Fax: +61-3-8344-5037, E-mail:

Copyright © 2016, Korean Society for Food Science of Animal Resources. This is an open access article distributed under the terms of the Creative Commons Attribution Non-Commercial License ( which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

Received: Sep 14, 2016 ; Revised: Oct 08, 2016 ; Accepted: Oct 17, 2016

Published Online: Dec 31, 2016


Fat in meat greatly improves eating quality, yet many consumers avoid visible fat, mainly because of health concerns. Generations of consumers, especially in the English-speaking world, have been convinced by health authorities that animal fat, particularly saturated or solid fat, should be reduced or avoided to maintain a healthy diet. Decades of negative messages regarding animal fats has resulted in general avoidance of fatty cuts of meat. Paradoxically, low fat or lean meat tends to have poor eating quality and flavor and low consumer acceptability. The failure of low-fat high-carbohydrate diets to curb “globesity” has prompted many experts to re-evaluate of the place of fat in human diets, including animal fat. Attitudes towards fat vary dramatically between and within cultures. Previous generations of humans sought out fatty cuts of meat for their superior sensory properties. Many consumers in East and Southeast Asia have traditionally valued more fatty meat cuts. As nutritional messages around dietary fat change, there is evidence that attitudes towards animal fat are changing and many consumers are rediscovering and embracing fattier cuts of meat, including marbled beef. The present work provides a short overview of the unique sensory characteristics of marbled beef and changing consumer preferences for fat in meat in general.

Keywords: intramuscular fat; animal fat; fatty meat cuts; marbling; marbled beef; meat quality


The fat in meat plays an important role in overall meat palatability. Intramuscular fat (IMF) positively influences sensory quality traits of meat including flavor, juiciness and tenderness, whereas a low amount of fat induces a less positive response (Hocquette et al., 2010; Van Elswyk and McNeill, 2014). The amount of visible fat, both subcutaneous and intramuscular, is used as a visual cue by consumers to judge meat quality. Many Asian consumers prefer moderate amounts of IMF in the meat they purchase, whereas visible fat is unpopular with Western (e.g., Europe, Australia) consumers. The amount of IMF within beef muscle varies widely depending on animal breed or genetics (Barendse, 2014; Hocquette et al., 2014; Widmann et al., 2011), the nutrition system (pasture or grain) (Duckett et al., 2013; Sithyphone et al., 2011; Van Elswyk and McNeill, 2014), animal maturity and weight (Duckett et al., 2014; Frank et al., 2016), primal (Pavan and Duckett, 2013) and other factors. Marbling describes the small flecks of visible fat deposited between individual muscle fiber bundles. Increases in IMF are generally accompanied by improvements in tenderness, juiciness and mouthfeel (Thompson, 2004). Therefore, this review addresses the importance of fat in beef for sensory palatability and consumer satisfaction.

The Importance of Fat in Meat for Consumer Satisfaction

Grilled beef flavor is due to a combination of heat generated aroma volatiles and non-volatile taste compounds (mainly free amino acids, peptides and organic acids) delivered in a unique matrix of muscle fiber, collagen, warmed-meat juices and partly dissolved fat (Frank et al., 2016). The positive relationship between IMF and overall palatability (tenderness and juiciness) and liking has been firmly established, mainly by untrained or naive consumer panels (Corbin et al., 2015; Emerson et al, 2013; Hunt et al., 2014; O'Quinn et al., 2012; Thompson, 2004). There have been fewer studies regarding the impacts of IMF on beef flavor using trained sensory panels (Frank et al., 2016; Legako et al., 2016). Trained panels learn to objectively rate the intensity of defined sensory attributes using validated quantitative scales. Consumers are generally asked to provide hedonic information about liking of products, whereas trained panels are used to provide objective quantitative information. The quality of data obtained using trained panels is more accurate than data obtained from untrained consumers. In one recent study comparing low and high fat Wagyu, a trained sensory panel found no difference in either tenderness or flavor, although the high fat samples were rated higher for juiciness (Okumura et al., 2007). In contrast, a positive curvilinear relationship between IMF and beef flavor scores was reported over a range of 0.3% to 15% fat, that plateaued at higher IMF content (Thompson, 2004). More recent studies confirm a strong correlation between marbling level and flavor (Corbin et al., 2015; Frank et al., 2016; Jung et al., 2016; Legako et al., 2015; Mateescu et al., 2015; O'Quinn et al., 2012).

Some Reasons Why We Like Fatty Meat

Fat is an essential macronutrient and a vehicle for fat soluble vitamins. Fat also plays a critical role in defining the sensory properties of complex foods, such as marbled beef. Apart from making food softer, fat facilitates “oral processing”, lubrication of food particles, increases saliva viscosity and acts as a binder or glue assisting in formation of a solid bolus in preparation for swallowing (Almiron-Roig, 2010; de Lavergne et al., 2015; Foster et al., 2011; Frank et al., 2015; Salles et al., 2011). There is increasing evidence for the existence of a long chain fatty acid taste receptor mechanism (Galindo et al., 2012; Tucker et al., 2014; Voigt et al., 2014) and human preferences for fat have been linked to variations in saliva flow and composition (Méjean et al., 2015; Mennella et al., 2014; Running et al., 2013). Fat may also increase parasympathetic saliva production, perhaps through the influence of free fatty acids, increasing perceived juiciness, although supporting data is lacking (Hodson and Linden, 2004).

Fat produces “mouth-coating” sensations, where a film of fat remains on oral membranes; this may result in greater persistence of fat soluble flavors in the mouth (Frank et al., 2011). As the level of marbling increases in meat, the bulk density decreases; there is less muscle fiber and collagen per unit volume of meat, potentially requiring less oral processing (Foster et al., 2011; Frank, 2015; Salles et al., 2011). Increasing levels of IMF may also contribute to meat tenderization by disrupting the organization of intramuscular connective tissue (Li et al., 2006). Fat plays an important functional role in meat; fat conducts heat more slowly than water, meat with a high fat content takes longer to reach a desired internal temperature, compared to meat with a low fat content (Baghekhandan et al., 1982). In practical terms this means that when cooking to the same endpoint temperature, meat of high fat content will have longer exposure to grilling surfaces and greater potential for generation of aroma volatiles and taste compounds (Frank et al., 2016). It should be noted that it is well-known that other factors make a contribution to meat tenderness such as the amount of collagen, and protein, cross-linking (Archile-Contreras et al., 2010; Purslow, 2014; Starkey et al., 2015).

Fat and Beef Flavor

Fat acts both as a substrate and a reservoir for flavor compounds and also affects the temporal flavor release. The role of unsaturated lipids and phospholipids in the formation of volatile compounds is well known (Elmore et al., 2000; Elmore et al., 2002). Different types of lipids have been shown to either promote or suppress the formation of different classes of volatiles in the presence of various amino acids (Farmer and Mottram, 1990; Farmer and Mottram, 1992; Farmer et al., 1989). Highly reactive radical intermediates from lipid oxidation can interact with other molecules to produce an array of potent aroma molecules formed through the Strecker degradation and other complex reactions (Frankel, 1980; Hidalgo et al., 2013; Zamora et al., 2015). There are many inconsistencies in the literature, past and present, regarding which lipid substrates contribute to volatile formation in meat, though it is widely believed that unsaturated free fatty acids and phospholipids contribute the most. Under the extreme temperature conditions of grilling and frying, many complex reactions take place leading not only to generation of aroma volatiles, but also reactions on the meat surface where tasteactive polymeric condensation products, known as melonoidin, formed - Melonoidin is also responsible for the desirable browned surface of grilled meat (Adams et al., 2009; Obretenov et al., 1993). Unfortunately, mutagenic heterocyclic aromatic amines (HAAs) are also generated at high grilling temperatures. Paradoxically, the level of HAAs is positively correlated to flavor and liking (Gibis et al., 2015). Recent studies show that beef with higher fat content produces lower levels of HAAs, especially fat from grass fed animals (Szterk et al., 2014).

In a recent study on grilled beef from three different production types, grass-fed Wagyu, grass-fed Angus and grain-fed Angus, a trained sensory panel showed that most flavor-related attributes were significantly correlated with the level of IMF (Frank et al., 2016). For example, apart from the expected improvements in juiciness and tenderness, as the level of marbling increased the meat became sweeter and saltier and the acidity, astringency and after-taste decreased. The odor and flavor impact, grilled beef, bloody, dairy fat and grassy flavors increased with marbling level, whereas hay/grainy and livery flavors decreased. Some of these relationships are shown in Fig. 1. These trends were consistent regardless of the breed or feed, implying a degree of generalization can be used for describing sensory properties of marbled beef. In the same study, it was shown that the concentration of most aroma-active volatiles increased with the marbling level (Table 1). The greatest amount of volatiles was measured in grass-fed Wagyu compared to grain-fed Angus, corresponding with higher flavor scores in the Wagyu samples. The concentration of non-volatile flavor compounds, such as free amino acids and carnosine also increased in beef after grilling, especially in high IMF Wagyu. Organic acids such as succinic and α-ketoglutarate, positively associated with meat flavor, increased with IMF, and lactic acid, associated with sourness, decreased (Table 1). In an other recent study on Hanwoo beef, the significant correlation between high IMF and positive sensory palatability attributes was confirmed (Jung et al., 2015; Jung et al., 2016). It was suggested that the palatability of Hanwoo beef can be improved by increasing IMF content, as increased IMF content leads to an increase in sensory tenderness, flavor, and juiciness.

Fig. 1. Effect of increasing intramuscular fat (IMF) on trained panel sensory scores for various attributes for various attributes (Damian Frank, unpublished results).
Download Original Figure
Table 1. Effect of different levels of intramuscular fat (low, medium, high) in beef on water-holding capacity characteristics, content of amino acids important for flavor and trained panel assessments. The average IMF (%) for the group is shown in brackets for each trait and level of IMF
Low IMF Medium IMF High IMF
Flavor - chemical and sensory
Total umami/ sweet amino acids (mg/kg)1 70a (7.63) 94b (12.65)
Aspartic acid (mg/kg)1 (an amino acid contributing to umami) 0.80a (7.63) 1.14b (12.65)
Trained panel - overall flavor impact (scale 0=100)1 60.39a (7.8) 60.87a (10.9) 63.95b (17.5)
Juiciness/water - chemical and sensory aspects
Moisture (%)3 73.68a (6.6) 69.45b (11.02) 60.93c (21.48)
Drip loss (%)2 6.26a (6.13) 4.53b (9.87)
Cook loss (%)1 27.0b (5.2) 25.8b (10.2) 23.6a (17.5)
Trained panel juiciness after 3 chews (scale 0=100)1 41.12b (7.8) 43.5b (10.9) 53.08a (17.5)
Trained panel juiciness after 10 chews (scale 0=100)1 36.24b (7.8) 38.4b (10.9) 46.82a (17.5)

a,bMeans with different superscripts within a row are significantly different (p<0.05).

1Frank et al., 2016; 2Kim and Lee, 2003; 3Jo et al., 2012.

Download Excel Table

Once formed, fat-soluble flavors naturally partition into the fat phase (e.g., melted fat) affecting their flavor release during eating and subsequent perception (Frank et al., 2011, 2012; Martuscelli et al., 2008). The fat solubility of aroma volatiles can be described by octanol water partitioning coefficients - LogP values; higher values indicate greater fat solubility. It was shown that aroma volatiles with high LogP values, such as nonanal and (E)-2-non-enal were less released into the headspace as IMF increased, whereas less fat soluble aroma volatiles increased, e.g.; 3-methylbutanal, 2,3-butanedione and 2-ethyl-3,5-dimethylpyrazine (Frank et al., 2016). The difference in ratios of aroma molecules at different IMF contents may explain some of the clear sensory changes perceived by consumers as the IMF increases. Fat also has a concentrating effect on water-soluble flavor molecules. For example sodium ions and many free amino acids do not readily dissolve in fat. By increasing the amount of fat in a food matrix, hydrophilic water soluble molecules are “pushed” into the aqueous phase increasing their relative concentration. This may increase the perceived intensity of the flavor (Chabanet et al., 2013; Lawrence et al., 2012). Similarly volatiles with low fat solubility (lipophobic molecules) may be released faster in the presence of significant levels of fat (Frank, 2015). Finally, fat has been shown to affect the rate of volatile release during eating in meat (Carrapiso, 2007; Lorido et al., 2015). In an unpublished study, we showed that the rate of release of an important water soluble beef aroma volatile (3-methylbutanal) was released faster, and at a higher concentration, in high fat Wagyu beef compared to low fat Wagyu beef (see Fig. 2) and was linked to positive differences in sensory perception (Fig. 1).

Fig. 2. In vivo measurement, using PTR-MS, of grilled beef volatiles during eating and swallowing of beef from two breeds (Angus, A; Wagyu, W), two feeding systems (Grass, G; Grain, GRN) and from two levels of marbling (High, H; L Low) respectively, for the acronyms in the legend. Each line is the average of 30 replicates. The black bars indicate the least significant difference at a time point. B = background breath, C1 = chew 1, etc. Extracted from Frank et al. (2014), Final report to MLA.
Download Original Figure

Quality Characteristics of High IMF Beef

Meat Standards Australia (MSA) uses a marbling score system (MSA-MB), based on the scheme developed by the United States Department of Agriculture (USDA). The MSA-MB system provides a fine scale for accurate measurement of beef marbling; the scoring system ranges from 100 to 1190 in increments of 10. Highly marbled meat is considered a premium product in Japan and Korea (Cho et al., 2010; Thompson, 2004). In Europe and Australia, increasing levels of marbling correspond to more acceptable flavor, juiciness and overall liking (Fig. 1).

The water-holding capacity and chemical composition of meat are influenced by intramuscular fat (Table 1). As the intramuscular fat increases from 5% to 22%, the moisture content declines (Jo et al., 2012). Consequently the drip loss and water lost during cooking are lower in meat with high IMF (Frank et al., 2016; Kim and Lee, 2003). As shown in Table 1 and in Fig. 1, a higher fat content in the meat also results in higher initial, and sustained, juiciness scores given by consumers and trained panelists. The reduced loss in water during storage and cooking is an advantage, as there is lower weight loss, improved yields, reduced loss of nutrients and greater returns to the wholesaler, retailer and food service industry.

The taste of meat is derived from a variety of compounds, including amino acids which can impart sweet, umami, salty, sour and other tastes (Frank et al., 2016). The amino acids contributing to sweet and umami taste have been found to be higher in beef striploin of higher IMF (Table 1). It is likely that these amino acids, in part, contribute to the increased consumer ‘like flavor’ scores (Fig. 3) and increased trained panel scores for ‘overall flavor impact’ (Table 2).

Fig. 3. Effect of muscle intramuscular fat (%) on predicted consumer scores for (a) flavor and (b) juiciness of grilled striploins, adjusted to a standard peak shear force (5.0 kg) and standard animal age (716 d). Vertical bars indicate standard error. From Thompson (2004).
Download Original Figure
Table 2. Effect of finishing system (pasture vs feedlot) and level of marbling on trained panel scores for flavor and ‘blind’ and ‘informed’ consumer scores for acceptability (n=8 loins per treatment). From Morales et al. (2013)
Pasture Feedlot p-value

Low marbling High marbling Low marbling High marbling
Trained panel Flavor1 4.5b 4.9a 4.4b 5.4a 0.031
Consumer - Blind accceptability2 4.9ab 4.9ab 4.6b 5.2a <0.001
Consumer - Informed acceptability3 5.4a 5.3a 5.0b 5.2ab 0.003

1Flavor; 0 = absence to 10 = maximum intensity.

2Each consumer received the four beef samples from each treatment, asked to evaluate acceptability; 1 = dislike extremely, 7 = like extremely.

3Each consumer received the four beef samples for each treatment, except each sample was accompanied by cards showing a photo corresponding to how the animal was raised (pasture vs feedlot) and the level of marbling in the raw meat (low vs high). The consumer gave an acceptability score as per the ‘blind’ acceptability.

Download Excel Table

Cultural Differences in Liking of Meat Characteristics

Many consumers prefer the appearance of beef with low or zero levels of marbling (Morales et al., 2013). But as shown in Table 1, when these same consumers are given cooked beef “blind”, without knowledge of the marbling level, they prefer the flavor of highly marbled beef and find it more acceptable (Morales et al., 2013). Consumers from Asian countries (Korea, Taiwan, Japan) generally prefer raw meat with a moderate amount of marbling (Ngapo et al., 2007).

In trials conducted in Japan, Korea and Australia using Meat Standards Australia (MSA) protocols, with beef ranging from low to high marbling, it was found that consumers from all three countries clearly identify eating quality differences and the grade cut-off scores given by Japanese and Korean consumers (fail, 3-star, 4-star, 5-star; corresponding to unsatisfactory, everyday, good everyday, premium, respectively) were very similar to the cut-offs given by Australian consumers (Park et al., 2008; Thompson et al., 2008). The weightings given in the meat quality score by Japanese and Korean consumers were more influenced by flavour and juiciness, than for Australian consumers. Korean consumers rated juiciness and flavor lower than Australian consumers across a range of beef muscles (Fig. 4). This may suggest that while tenderness is universally well understood, other attributes such as juiciness and flavor, may have specific cultural contexts.

Fig. 4. Effect of consumer (Korea vs Australia) and muscle (BB, Biceps brachii; LL, Longissimus lumborum; SM, semimembranosus) on the consumer scores for tenderness, juiciness and flavor (scale; 0 = lowest liking to 100 = highest liking). Standard error = 1.4, 1.1, 1.0 for tenderness, juiciness and flavor respectively. From Park et al., 2008.
Download Original Figure

Changes in Attitudes towards Animal Fat

It is very clear that consumer attitudes toward animal fats, especially the saturated fat in meat has been overwhelmingly negative in the US, Australia and other English speaking countries for the last 50 years (Ngapo and Dransfield, 2006; Williams and Droulez, 2010; Williams and Mummery, 2013). The per capita decline in beef consumption in the US and other Western countries has been attributed in large part to animal fat phobia (Higgs, 2000). Most public health authorities, continue to recommend reduction of saturated fat in the diet, including avoiding fatty cuts of meat. These health recommendations are obviously in conflict with the hedonic reality of a high fat product, such as marbled beef. A growing body of research questions the validity of the so called “diet-heart hypothesis” (Ramsden et al., 2016; Siri-Tarino et al., 2010; Smith et al., 2015; Willett, 2003) and calls for a re-evaluation of role of animal fats in the diet (Barendse, 2014; Klurfeld, 2015). Although a nutritional consensus on saturated fat is unlikely to be found soon, the overall position of fat in the modern diet appears to be changing. The failure of low fat, high carbohydrate diets to curb obesity is being acknowledged by health experts and in popular media (Drewnowski, 2015; Teicholz, 2014). Hence, inclusion of high fat foods with superior sensory properties in a balanced diet, is likely to gain wider acceptance in the near future.

Opportunities for Improving Meat Quality

An unfavorable balance of omega-6/omega-3 fatty acids in the Western diet has been associated with negative health impacts (Lawrence, 2013; Simopoulos, 2008). The balance has been shifting towards omega-6 fats partly as a consequence of industrial animal production (Barendse, 2014; Yu et al., 2013). Ruminants raised on less intensive pasture based systems generally have a more favorable ratio than animals raised on grain or concentrate. Beef can be a good source of mono-unsaturated oleic acid as well as short and long chain omega-3 fatty acids, with proven health benefits. Ruminants also produce a number of unique unsaturated fatty acids (vaccenic and rumenic) which probably have positive human health effects (Bjorklund et al., 2014; Daley et al., 2010; Pavan and Duckett, 2013). Importantly, unsaturated fat has been shown to significantly increase satiety compared to saturated fats (Maljaars et al., 2009). Smaller, more satiating portions of marbled beef high in “healthy” unsaturated fat, can play an important role in sustainable beef production.

Previous studies showed lower consumer acceptance of grass-fed beef compared to beef raised on concentrate (Bjorklund et al., 2014; Hunt et al., 2014; Legako et al., 2015; Maughan et al., 2012). Most comparisons are typically between very low fat grass-fed beef with much higher fat grain-fed produce (Van Elswyk and McNeill, 2014). The lower preference for low fat grass-fed meat is expected, given the clear impacts of IMF on beef sensory properties. Yet, when grass and grain fed marbled beef with the same IMF was compared, few distinctive grass-fed flavors were apparent, especially at levels of IMF above 5%. The data strongly suggested that differences in the flavor of grass and grain fed beef is mainly due to differences in fat level rather than due to inherent grass or grain flavors (Frank et al., 2016). Refinement of animal production and feeding strategies to optimize the composition of IMF and enhance sensory properties of beef will present new opportunities for producers of marbled beef (Barendse, 2014; Scollan et al., 2014; Widmann et al., 2011).


Fat in meat positively influences sensory quality traits of meat, and the positive relationship between IMF and overall palatability has been firmly established by both untrained and consumer panels. IMF plays an important functional role in meat and acts both as a substrate and a reservoir for flavor compounds. Research has shown that most flavor-related attributes are significantly correlated with the level of IMF. In this regard, consumers from Asian countries including Korea, Japan and Taiwan generally prefer meat with a moderate amount of marbling. Also, the overall position of fat in the modern diet appears to be changing, although a nutritional consensus on saturated fat is unlikely to be found soon. The failure of low fat, high carbohydrate diets to curb obesity is being acknowledged by health experts and in popular media. Beef can be a good source of oleic acid and short and long chain omega-3 fatty acids, with proven health benefits. Consequently, refinement of animal production and feeding strategies to optimize the composition of IMF and enhance sensory properties of beef will present new opportunities for producers of marbled beef.



Adams A., Kitryte V., Venskutonis R., De Kimpe N. Formation and characterisation of melanoidin-like polycondensation products from amino acids and lipid oxidation products. Food Chem. 2009; 115:904-911.


Almiron-Roig D. Human perceptipons and preferences for fat-rich foods. CRC Press. Boca Raton (FL), USA: 2010.


Archile-Contreras A. C., Mandell I. B., Purslow P. P. Disparity of dietary effects on collagen characteristics and toughness between two beef muscles. Meat Sci. 2010; 86:491-497.


Baghekhandan M. S., Okos M. R., Sweat V. E. The thermal-conductivity of beef as affected by temperature and composition. T. Asae. 1982; 25:1118-1122.


Barendse W. Should animal fats be back on the table? A critical review of the human health effects of animal fat. Anim. Prod. Sci. 2014; 54:831-855.


Bjorklund E. A., Heins B. J., DiCostanzo A., Chester-Jones H. Fatty acid profiles, meat quality, and sensory attributes of organic versus conventional dairy beef steers. J. Dairy Sci. 2014; 97:1828-1834.


Carrapiso A. I. Effect of fat content on flavour release from sausages. Food Chem. 2007; 103:396-403.


Chabanet C., Tarrega A., Septier C., Siret F., Salles C. Fat and salt contents affect the in-mouth temporal sodium release and saltiness perception of chicken sausages. Meat Sci. 2013; 94:253-261.


Cho S. H., Kim J., Park B. Y., Seong P. N., Kang G. H., Kim J. H., Jung S. G., Im S. K., Kim D. H. Assessment of meat quality properties and development of a palatability prediction model for Korean Hanwoo steer beef. Meat Sci. 2010; 86:236-242.


Corbin C. H., O'Quinn T. G., Garmyn A. J., Legako J. F., Hunt M. R., Dinh T. T. N., Rathmann R. J., Miller M. F. Sensory evaluation of tender beef strip loin steaks of varying marbling levels and quality treatments. Meat Sci. 2015; 100:24-31.


Daley C. A., Abbott A., Doyle P. S., Nader G. A., Larson S. A review of fatty acid profiles and antioxidant content in grass-fed and grain-fed beef. Nutr. J. 2010; 9:10.


de Lavergne M. D., van de Velde F., van Boekel M., Stieger M. Dynamic texture perception and oral processing of semi-solid food gels: Part 2: Impact of breakdown behaviour on bolus properties and dynamic texture perception. Food Hydrocol. 2015; 49:61-72.


Drewnowski A. The carbohydrate-fat problem: Can we construct a healthy diet based on dietary guidelines?. Adv. Nutr. 2015; 6:318S-325S.


Duckett S. K., Neel J. P. S., Lewis R. M., Fontenot J. P., Clapham W. M. Effects of forage species or concentrate finishing on animal performance, carcass and meat quality. J. Anim. Sci. 2013; 91:1454-1467.


Duckett S. K., Rosso C. F., Lagreca G. V., Miller M. C., Neel J. P. S., Lewis R. M., Swecker W. S., Fontenot J. P. Effect of frame size and time-on-pasture on steer performance, longissimus muscle fatty acid composition, and tenderness in a forage-finishing system. J. Anim. Sci. 2014; 92:4767-4774.


Elmore J. S., Campo M. M., Enser M., Mottram D. S. Effect of lipid composition on meat-like model systems containing cysteine, ribose, and polyunsaturated fatty acids. J. Agr. Food Chem. 2002; 50:1126-1132.


Elmore J. S., Mottram D. S., Enser M., Wood J. D. The effect of dietary polyunsaturated fatty acid supplements on the volatile composition of cooked sheep meat. Meat Sci. 2000; 55:149-159.


Emerson M. R., Woerner D. R., Belk K. E., Tatum J. D. Effectiveness of USDA instrument-based marbling measurements for categorizing beef carcasses according to differences in longissimus muscle sensory attributes. J. Anim. Sci. 2013; 91:1024-1034.


Farmer L. J., Mottram D. S. Interaction of lipid in the Maillard reaction between cysteine and ribose - the effect of tryglyceride and 3 phospholipids on the volatile products. J. Sci. Food Agr. 1990; 53:505-525.


Farmer L. J., Mottram D. S. Effect of cysteine and ribose on the volatile thermal-degradation products of a triglyceride and 3 phospholipids. J. Sci. Food Agr. 1992; 60:489-497.


Farmer L. J., Mottram D. S., Whitfield F. B. Volatile compounds produced in Maillard reactions involving cysteine, ribose and phospholipid. J. Sci. Food Agr. 1989; 49:347-368.


Foster K. D., Grigor J. M. V., Cheong J. N., Yoo M. J. Y., Bronlund J. E., Morgenstern M. P. The role of oral processing in dynamic sensory perception. J. Food Sci. 2011; 76:R49-R61.


Frank D., Appelqvist I., Piyasiri U., Delahunty C. In vitro measurement of volatile release in model lipid emulsions using proton transfer reaction mass spectrometry. J. Agr. Food Chem. 2012; 60:2264-2273.


Frank D., Appelqvist I., Piyasiri U., Wooster T., Delahunty C. Proton transfer reaction mass spectrometry and time intensity perceptual measurement of flavor release from lipid emulsions using trained human subjects. J. Agr. Food Chem. 2011; 59:4891-4903.


Frank D., Ball A., Hughes J., Krishnamurthy R., Piyasiri U., Stark J., Watkins P., Warner R. Sensory and flavor chemistry characteristics of Australian beef: Influence of intramuscular fat, feed and breed. J. Agr. Food Chem. 2016; 64:4299-4311.


Frank D., Eyres G. T., Piyasiri U., Cochet-Broch M., Delahunty C. M., Lundin L., Appelqvist I. M. Effects of agar gel strength and fat on oral breakdown, volatile release and sensory perception using in vivo and in vitro systems. J. Agr. Food Chem. 2015; 63:9093-9102.


Frankel E. N. Lipid oxidation. Prog. Lip. Res. 1980; 19:1-22.


Galindo M. M., Voigt N., Stein J., van Lengerich J., Raguse J. D., Hofmann T., Behrens M. G protein-coupled receptors in human fat taste perception. Chem. Senses. 2012; 37:123-139.


Gibis M., Kruwinnus M., Weiss J. Impact of different pan-frying conditions on the formation of heterocyclic aromatic amines and sensory quality in fried bacon. Food Chem. 2015; 168:383-389.


Hidalgo F. J., Alcon E., Zamora R. Cysteine- and serine-thermal degradation products promote the formation of Strecker aldehydes in amino acid reaction mixtures. Food Res. Int. 2013; 54:1394-1399.


Higgs J. D. The changing nature of red meat: 20 years of improving nutritional quality. Trends Food Sci. Tech. 2000; 11:85-95.


Hocquette J. F., Gondret F., Baza E. M., dale F., Jurie C., Pethick D. W. Intramuscular fat content in meat-producing animals: Development genetic and nutritional control and identification of putative markers. Animal. 2010; 4:303-319.


Hocquette J. F., Van Wezemael L., Chriki S., Legrand I., Verbeke W., Farmer L., Scollan N. D., Polkinghorne R., Rodbotten R., Allen P., Pethick D. W. Modelling of beef sensory quality for a better prediction of palatability. Meat Sci. 2014; 97:316-322.


Hodson N. A., Linden R. W. Is there a parotid-salivary reflex response to fat stimulation in humans?. Physiol. Behav. 2004; 82:805-813.


Hunt M. R., Garmyn A. J., O'Quinn T. G., Corbin C. H., Legako J. F., Rathmann R. J., Brooks J. C., Miller M. F. Consumer assessment of beef palatability from four beef muscles from USDA choice and select graded carcasses. Meat Sci. 2014; 98:1-8.


Jo C., Cho S. H., Chang J., Nam K. C. Keys to production and processing of Hanwoo beef: A perspective of tradition and science. Anim. Front. 2012; 2:32-38.


Jung E.Y., Hwang Y. H., Joo S. T. Chemical components and meat quality traits related to palatability of ten primal cuts from Hanwoo carcasses. Kor. J. Food Sci. An. 2015; 35:859-866.


Jung E. Y., Hwang Y. H., Joo S. T. The relationship between chemical composition, meat quality, and palatability of the 10 primal cuts from Hanwoo steer. Kor. J. Food Sci. An. 2016; 36:137-143.


Kim C. J., Lee E. S. Effects of quality grade on the chemical, physical and sensory characteristics of Hanwoo (Korean native cattle) beef. Meat Sci. 2003; 63:397-405.


Klurfeld D. M. Research gaps in evaluating the relationship of meat and health. Meat Sci. 2015; 109:86-95.


Lawrence G., Buchin S., Achilleos C., Berodier F., Septier C., Courcoux P., Salles C. In vivo sodium release and saltiness perception in solid lipoprotein matrices. 1. Effect of composition and texture. J. Agr. Food Chem. 2012; 60:5287-5298.


Lawrence G. D. Dietary fats and health: Dietary recommendations in the context of scientific evidence. Adv. Nutr. 2013; 4:294-302.


Legako J. F., Dinh T. T. N., Millere M. F., Adhikari K., Brooks J. C. Effects of USDA quality grade on consumer palatability scores, trained descriptive attributes and volatile compounds of grilled beef. Meat Sci. 2015; 100:246-255.


Legako J. F., Dinh T. T. N., Miller M. F., Adhikari K., Brooks J. C. Consumer palatability scores, sensory descriptive attributes, and volatile compounds of grilled beef steaks from three USDA quality grades. Meat Sci. 2016; 112:77-85.


Li C., Zhou G., Xu X., Zhang J., Xu S., Ji Y. Effects of marbling on meat quality characteristics and intramuscular connective tissue of beef Longissimus muscle. Asian-Aust. J. Anim. Sci. 2006; 19:1799-1808.


Lorido L., Estevez M., Ventanas J., Ventanas S. Salt and intramuscular fat modulate dynamic perception of flavour and texture in dry-cured hams. Meat Sci. 2015; 107:39-48.


Maljaars J., Romeyn E. A., Haddeman E., Peters H. P., Masclee A. A. Effect of fat saturation on satiety, hormone release, and food intake. Am. J. Clin. Nutr. 2009; 89:1019-1024.


Martuscelli M., Savary G., Pittia P., Cayot N. Vapour partition of aroma compounds in strawberry flavoured custard cream and effect of fat content. Food Chem. 2008; 108:1200-1207.


Mateescu R. G., Garrick D. J., Garmyn A. J., VanOverbeke D. L., Mafi G. G., Reecy J. M. Genetic parameters for sensory traits in longissimus muscle and their associations with tenderness, marbling score, and intramuscular fat in Angus cattle. J. Anim. Sci. 2015; 93:21-27.


Maughan C., Tansawat R., Cornforth D., Ward R., Martini S. Development of a beef flavor lexicon and its application to compare the flavor profile and consumer acceptance of rib steaks from grass- or grain-fed cattle. Meat Sci. 2012; 90:116-121.


Méjean C., Morzel M., Neyraud E., Issanchou S., Martin C., Bozonnet S., Urbano C., Schlich P., Hercberg S., Péneau S, Feron G. Salivary composition is associated with liking and usual nutrient intake. PLoS ONE. 2015; 10:e0137473.


Mennella I., Fogliano V., Vitaglione P. Salivary lipase and et-amylase activities are higher in overweight than in normal weight subjects: Influences on dietary behavior. Food Res. Int. 2014; 66:463-468.


Morales R., Aguiar A. P. S., Subiabre I., Realini C. E. Beef acceptability and consumer expectations associated with production systems and marbling. Food Qual. Prefer. 2013; 29:166-173.


Ngapo T. M., Dransfield E. British consumers preferred fatness levels in beef: Surveys from 1955, 1982 and 2002. Food Qual. Prefer. 2006; 17:412-417.


Ngapo T. M., Martin J. F., Dransfield E. International preferences for pork appearance: II. Factors influencing consumer choice. Food Qual. Prefer. 2007; 18:139-151.


O'Quinn T. G., Brooks J. C., Polkinghorne R. J., Garmyn A. J., Johnson B. J., Starkey J. D., Rathmann R. J., Miller M. F. Consumer assessment of beef strip loin steaks of varying fat levels. J. Anim. Sci. 2012; 90:626-634.


Obretenov T. D., Ivanova S. D., Kuntcheva M. J., Somov G. T. Melanoidin formation in cooked meat products. J. Agr. Food Chem. 1993; 41:653-656.


Okumura T., Saito K., Nade T., Misumi S., Masuda Y., Sakuma H., Kawamura T. Effects of intramuscular fat on the sensory characteristics of M-longissimus dorsi in Japanese black steers as judged by a trained analytical panel. Asian-Aust. J. Anim. Sci. 2007; 20:577-581.


Park B. Y., Hwang I. H., Cho S. H., Yoo Y. M., Kim J. H., Lee J. M., Polkinghorne R., Thompson J. M. Effect of carcass suspension and cooking method on the palatability of three beef muscles as assessed by Korean and Australian consumers. Aust. J. Exp. Agr. 2008; 48:1396-1404.


Pavan E., Duckett S. K. Fatty acid composition and interrelationships among eight retail cuts of grass-feed beef. Meat Sci. 2013; 93:371-377.


Purslow P. P. New developments on the role of intramuscular connective tissue in meat toughness. Ann. Rev. Food Sci. Tech. 2014; 5:133-153.


Ramsden C. E., Zamora D., Majchrzak-Hong S., Faurot K. R., Broste S. K., Frantz R. P., Davis J. M., Ringel A., Suchindran C. M., Hibbeln J. R. Re-evaluation of the traditional diet-heart hypothesis: Analysis of recovered data from Minnesota Coronary Experiment (1968-73). BMJ. 2016; 353i1246.


Running C. A., Mattes R. D., Tucker R. M. Fat taste in humans: Sources of within- and between-subject variability. Prog. Lipid Res. 2013; 52:438-445.


Salles C., Chagnon M. C., Feron G., Guichard E., Laboure H., Morzel M., Semon E., Tarrega A., Yven C. In-mouth mechanisms leading to flavor release and perception. Crit. Rev. Food Sci. Nutr. 2011; 51:67-90.


Scollan N. D., Dannenberger D., Nuernberg K., Richardson I., MacKintosh S., Hocquette J.-F., Moloney A. P. Enhancing the nutritional and health value of beef lipids and their relationship with meat quality. Meat Sci. 2014; 97:384-394.


Simopoulos A. P. The importance of the omega-6/omega-3 fatty acid ratio in cardiovascular disease and other chronic diseases. Exp. Biol. Med. 2008; 233:674-688.


Siri-Tarino P. W., Sun Q., Hu F. B., Krauss R. M. Meta-analysis of prospective cohort studies evaluating the association of saturated fat with cardiovascular disease. Am. J. Clin. Nutr. 2010; 91:535-546.


Sithyphone K., Yabe M., Horita H., Hayashi K., Fumita T., Shiotsuka Y., Etoh T., Ebara F., Samadmanivong O., Wegner J., Gotoh T. Comparison of feeding systems: feed cost, palatability and environmental impact among hayfattened beef, consistent grass-only-fed beef and conventional marbled beef in Wagyu (Japanese Black cattle). Anim. Sci. J. 2011; 82:352-359.


Smith J. D., Hou T., Ludwig D. S., Rimm E. B., Willett W., Hu F. B., Mozaffarian D. Changes in intake of protein foods, carbohydrate amount and quality, and long-term weight change: Results from 3 prospective cohorts. Am. J. Clin. Nutr. 2015; 101:1216-1224.


Starkey C. P., Geesink G. H., Oddy V. H., Hopkins D. L. Explaining the variation in lamb longissimus shear force across and within ageing periods using protein degradation, sarcomere length and collagen characteristics. Meat Sci. 2015; 105:32-37.


Szterk A., Waszkiewicz-Robak B. Influence of selected quality factors of beef on the profile and the quantity of heterocyclic aromatic amines during processing at high temperature. Meat Sci. 2014; 96:1177-1184.


Teicholz N. The Big Fat Surprise: Why butter, meat, and cheese belong in a healthy diet. Scribe Publications Pty Limited. 2014.


Thompson J. M. The effects of marbling on flavour and juiciness scores of cooked beef, after adjusting to a constant tenderness. Aust. J. Exp. Agr. 2004; 44:645-652.


Thompson J. M., Polkinghorne R., Hwang I. H., Gee A. M., Cho S. H., Park B. Y., Lee J. M. Beef quality grades as determined by Korean and Australian consumers. Aust. J. Exp. Agr. 2008; 48:1380-1386.


Tucker R. M., Mattes R. D., Running C. A. Mechanisms and effects of “fat taste” in humans. Biofactors. 2014; 40:313-326.


Van Elswyk M. E., McNeill S. H. Impact of grass/forage feeding versus grain finishing on beef nutrients and sensory quality: The US experience. Meat Sci. 2014; 96:535-540.


Voigt N., Stein J., Galindo M. M., Dunkel A., Raguse J. D., Meyerhof W., Hofmann T., Behrens M. The role of lipolysis in human orosensory fat perception. J. Lipid Res. 2014; 55:870-882.


Widmann P., Nuernberg K., Kuehn C., Weikard R. Association of an ACSL1 gene variant with polyunsaturated fatty acids in bovine skeletal muscle. BMC Genet. 2011; 12:96.


Willett W. C. Dietary fat and obesity: Lack of an important role. Scand. J. Nutr. 2003; 47:58-67.


Williams P., Droulez V. Australian red meat consumption - implications of changes over 20 years on nutrient composition. Food Aust. 2010; 62:87-94.


Williams S. L., Mummery K. W. Characteristics of consumers using 'better for you' front-of-pack food labelling schemes - an example from the Australian heart foundation tick. Public Health Nutr. 2013; 16:2265-2272.


Yu M., Gao Q., Wang Y., Zhang W., Li L., Wang Y., Dai Y. Unbalanced omega-6/omega-3 ratio in red meat products in China. J. Biomed. Res. 2013; 27:366-371.


Zamora R., Navarro J. L., Aguilar I., Hidalgo F. J. Lipid-derived aldehyde degradation under thermal conditions. Food Chem. 2015; 174:89-96.