PROTEIN: All of Your Burning Questions Answered By Alan Aragon - FIRST EDITION Copyright: ©2021 By Alan Aragon All rights reserved. This book or any part thereof, may not be reproduced or recorded in any form without permission in writing from the author, except for the quotation of brief excerpts, to which proper attribution is given. Suggested citation: Aragon AA. Protein: All of Your Burning Questions Answered. 1st ed., Los Angeles: Alan Aragon. https://alanaragon.com/books Cover art by Jeana Aragon (@jeana.aragon on Instagram) This book is NOT intended for the treatment or prevention of disease, nor as a substitute for medical treatment, nor as an alternative to medical advice. It is a review of scientific literature presented for informational purposes, to increase public knowledge of the developments in the field of nutrition. The information herein should not be adopted without consulting your doctor or qualified healthcare professional. Use of the information herein is at the sole choice and risk of the reader. The author and publisher specifically disclaim all responsibility for any liability, loss, or risk (personal or otherwise) incurred as a direct or indirect consequence of the application of any of the contents of this book. -1- This book is dedicated to Jeana, Lex, and Max. You are my dream team and my life’s joy. -2- Contents Chapter 1: Hierarchy of importance (start here, then skip through the book as you wish) [5] Chapter 2: How much protein does the general public habitually consume? [8] Chapter 3: How much protein is needed to maintain health in the general population? [10] Chapter 4: How much protein do competitive athletes habitually consume? [13] Chapter 5: How much protein maximizes athletic performance? [16] Chapter 6: How much protein maximizes muscle gain? [18] Chapter 7: How should protein be distributed through the day to maximize muscle gain? [24] Chapter 8: How much protein maximally preserves muscle while losing fat? [27] Chapter 9: How should protein be distributed through the day to maximize muscle retention while dieting? [30] Chapter 10: How does protein intake influence recomposition? [36] Chapter 11: How does protein timing (pre, during, and postexercise) impact athletic performance? [39] Chapter 12: How does protein timing (pre, during, and postexercise) impact body composition? [43] Chapter 13: How does BCAA supplementation affect body composition? [49] -3- Chapter 14: How does a high protein intake impact ketosis? [53] Chapter 15: How does protein restriction influence longevity? [56] Chapter 16: How does advanced age impact protein utilization and dosing requirements? [62] Chapter 17: How much protein do adolescents need? [67] Chapter 18: How do sex differences influence protein requirements? [69] Chapter 19: How do high-protein diets impact bone and kidney health? [73] Chapter 20: Is there an inherent advantage to pre-bed casein? [76] Chapter 21: How does postworkout whey compare to chicken, beef, or casein for improving body composition & strength? [79] Chapter 22: How do whole eggs compare with egg whites for muscle growth? [82] Chapter 23: If collagen is considered a low-quality protein, does that make collagen supplements useless? [85] Chapter 24: How do plant proteins compare to animal proteins for muscle growth? [87] Chapter 25: How satiating is protein, really? [92] Chapter 26: What about non-linear protein intake through the week (training days vs non-training days, protein hyperfeeds)? [95] Chapter 27: Protein servings & sources [101] Postscript [108] References [109] -4- Chapter 1: Hierarchy of importance (start here, then skip through the book as you wish) Welcome! Welcome, and thank you for cracking open this little beast. In the odd case that you’re reading this, yet you don’t know who I am or why I’m worth listening to, here is my bio, and here are my peer reviewed publications. The content of this book is meticulously compiled from the current state of the scientific evidence combined with nearly three decades of field experience. I’m proud to say that I’ve co-authored several of the key research publications that have shaped the current practice guidelines on protein intake for sports and fitnessoriented populations. The aim of this book is to provide a highly focused, fluff-free resource that concisely answers the most frequently asked protein questions I’ve encountered throughout my career as a trainer, nutritional counselor, researcher, and educator. So yes, get excited. Get very excited. :) Hierarchy of importance Here’s the crux of why I wanted you to read this introductory section first. Without maintaining the proper big-picture -5- perspective, the smaller details will lack meaning and context. As you go through the material, keep in mind that there’s an underlying order of importance when it comes to the various aspects about protein. From most to least important, the ranking is as follows (keep in mind that these are specific to protein; this hierarchy does not apply to all nutrients): 1) Total daily protein amount. For most of you reading this, getting total daily amount right is the most influential factor. Still, for a minority of individuals with very limited options, it’s theoretically possible to get total daily intake right, but lack quality (sufficient essential amino acids within bioavailable contexts1). We’ll be operating under the assumption that quality of the dietary protein sources is high overall. Therefore, from a practical standpoint, total daily amount is king. 2) Distribution of protein through the day – in other words, the spread or pattern of intake, including number of feedings and protein dose per meal. The impact of a more evenly spread versus skewed pattern, or a low vs. high feeding frequency, or a narrow versus broad feeding window depends on the individual goal. Nevertheless, these aspects -6- of within-day distribution are of distantly secondary importance compared to total daily amount. 3) Timing of protein relative to the training bout. For most goals, this factor has the least impact, especially in the context of programs with typical protein feeding distributions amounting to the proper daily total. Exceptions where protein timing relative to the training bout warrants attention are programs with very low meal frequency (e.g., 1-2 meals per day). In the latter case, the positioning/timing of protein can potentially influence rates of progress. The finer details of these concepts are elucidated in subsequent chapters. Caveats Keep in mind that nutritional needs vary across the stages of the human life cycle, well as different disease states. The protein requirements discussed in this book apply to healthy adults, unless specified otherwise. With that out of the way, let’s dive in! -7- Chapter 2: How much protein does the general public habitually consume? A common statement made by professors throughout my college experience was that protein supplements (and the push for greater protein intakes in fitness-related media) are a scam because people already consume more than enough protein. A more nuanced approach to this topic shows that different goals warrant different protein intakes. So, this claim always set off my skepticism sensors. The latest protein consumption data from the National Health and Nutrition Examination Survey (NHANES)2 shows that men aged 19-50 years consume 101.2-109.5 g/day. Using NHANES data, the Centers for Disease Control and Prevention (CDC) reported that the average bodyweight of men in the US is 89.8 kg.3 This amounts to an average protein intake of approximately 1.17 g/kg in men. NHANES data for women aged 19-50 years showed an intake of 70.3-72.9 g/day.2 The CDC reported an average bodyweight of 77.4 kg.3 This works out to a protein intake of approximately 0.92 g/kg in women. Both of these protein intakes (1.17 & 0.92 g/kg in men & women, respectively) exceed the Recommended Daily -8- Allowance (RDA) for protein, which is 0.8 g/kg.4 So, by that standard, my professors were correct. The general public’s protein intake exceeds the “official” public health guideline. The problem is that the RDA is insufficient to meet the needs of a substantial proportion of the general population, and it comes up short for practically all dieting and athletic populations. The following chapters cover the RDA’s shortcomings in more detail. -9- Chapter 3: How much protein is needed to maintain health in the general population? Ancient & tenacious The Recommended Daily Allowance (RDA) for protein is 0.8 g/kg.4 Notably, this figure was derived from nitrogen balance studies on sedentary individuals. It formally became part of the public health guidelines in 1980. It’s now 2021, and the RDA for protein hasn’t changed; no adjustments for athletes or physically active individuals, no increase for the elderly. Forty years is a long time for a guideline as important as protein intake to be outdated, despite a mountain of research showing benefits of greater intakes across virtually all populations. But, this is pretty much the normal (glacial) pace of conventional wisdom when it comes to altering established nutrition rules in general. Moving forward The research community’s call to re-evaluate the RDA has been ongoing and vigorous. A memorable 2009 review by Donald Layman,5 one of the pioneers of protein research in physically active subjects, was blatantly titled, Dietary Guidelines should reflect new understandings about adult protein needs. This was - 10 - perhaps the first paper to address the RDA’s lack of contingencies for protein needs based on the state of energy balance. Layman accurately contended that protein requirements are inversely proportional to energy intake. In other words, protein requirements increase in the face of hypocaloric (energy deficit) conditions, which pose an inherent threat to lean mass preservation. Furthermore, Layman noted beneficial effects on calcium metabolism and bone health at protein intakes above 1.2 g/kg. Protein requirements for the general adult population are largely focused on preserving lean mass in compromising conditions such as dieting and aging (separate goals from getting jacked or enhancing athletic performance, covered elsewhere in this book). A review by Lonnie et al6 relayed the collective guidelines of the International PROT-AGE Study Group and European Society for Clinical Nutrition and Metabolism (ESPEN) for individuals over age 65, which are as follows: 1.0-1.2 g/kg for healthy folks, 1.2-1.5 g/kg for those with acute or chronic illnesses, and 2.0 g/kg for those with severe illnesses, injuries, or malnutrition. A review by Phillips et al7 took direct aim at the RDA, focusing on protein needs for optimizing health and longevity, arriving - 11 - at a recommendation of 1.2-1.6 g/kg. This recommendation was inclusive of the general adult population, from younger to older. On the more generous end, Pencharz et al8 proposed an intake of 1.5-2.2 g/kg. Interestingly this recommendation pertained to the general population (not strength athletes or bodybuilders). However, their reasoning for this range was based, in part, on the Institute of Medicine’s Acceptable Daily Macronutrient Range (ADMR) of 10-35% of total energy intake,9 which carries a high degree of subjectivity. As is typical in research, the recommendations vary according to different perspectives and interpretations of the data. Among these recommendations, the intake range best supported by the research evidence for the healthy, notnecessarily-athletic-nor-dieting general public is Phillips et al’s proposed guideline of 1.2-1.6 g/kg.7 In Imperial terms, this translates to 0.54-0.72 g/lb. - 12 - Chapter 4: How much protein do high-level competitive athletes habitually consume? Strength/power athletes Gillen et al10 reported that elite-level Dutch strength athletes (71 subjects) had a protein intake that averaged 1.8 g/kg in those who used protein supplements, and 1.5 g/kg in those who did not. Slater and Phillips10 relayed the reported intakes of various strength/power athletes at the elite, national, and international levels as follows: Throwing: Sprinting: Weightlifting: Men 1.3-2.4 g/kg 1.5 g/kg 1.3-3.2 g/kg Women 1.1-2.5 g/kg 1.7 g/kg [no data] Bodybuilders Slater and Phillips11 reported that the protein intakes of elitelevel male and female bodybuilders was 1.7-2.4 g/kg & 1.5-2.0 g/kg, respectively. A systematic review by Spendlove et al12 reported a range of 157 g/day (1.9 g/kg/day) to 406 g/day (4.3 g/kg/day) among a mix of drug-free and enhanced competitive bodybuilders. Chappell et al13 reported that in high-level drugfree bodybuilders, pre-contest protein intakes of men and - 13 - women who placed in the top-5 were 3.3 & 2.8 g/kg, respectively. Protein intake of men and women who placed out of the top-5 were 2.7 & 2.9 g/kg, respectively. Body composition was not reported in this study, so no intakes based on fat-free mass can be reported. Mixed/team sports Team sports fall somewhere in the middle of the strengthendurance continuum, with a mix of demands and energy system contributions on that continuum. In a large sample of elite-level team sport athletes (242 subjects), Gillen et al10 found that protein intake averaged 1.6 g/kg in subjects who used protein supplements, and 1.4 g/kg in those who did not. A systematic review by Jenner et al14 reported the intakes of various professional & semi-professional mixed/team sports athletes as follows: Football (soccer): Australian football: Rugby union: Wheelchair basketball: Volleyball: Ice hockey: Men 1.9-2.0 g/kg 1.8-3.4 g/kg 2.2-2.7 g/kg 1.7 g/kg [no data] [no data] Women [no data] [no data] [no data] [no data] 0.9 g/kg 1.4 g/kg - 14 - Endurance athletes Finally, we have our quirky friends who just love to see how far they can push the limits of their fuel tanks. A classic review by Tarnopolsky et al15 reported protein intakes ranging 1.0-2.2 g/kg among high-level male and female endurance athletes. More recently, Burke et al16 reported that in elite-level Australian endurance athletes (4 canoeists, 2 cyclists, 11 distance runners, 3 kayakers, 9 rowers, 9 swimmers, and 3 walkers), protein intake averaged 1.9 g/kg. - 15 - Chapter 5: How much protein maximizes athletic performance? The following table outlines the latest position stands of the major nutrition (& exercise) organizations on protein requirements for athletic populations. The protein recommendation of the ISSN17 has been the same since their initial position stand on this topic in 2007; they were a bit ahead of the game. The current recommendation of the Academy of Nutrition and Dietetics, Dietitians of Canada, and American College of Sports Medicine18 has been increased since their previous statement in 2009, where the range was 1.2-1.7 g/kg. POSITION STANDS ON PROTEIN INTAKE FOR ATHLETIC GOALS Publication Population Recommendation Jäger R, et al. International Society of Sports Nutrition Position Stand: protein and exercise. J Int Soc Sports Nutr. 2017 Jun 20;14:20. [PubMed] Physically active individuals, including competitive and recreational athletes aiming to enhance muscular strength, endurance, or size 1.4-2.0 g/kg Thomas DT, et al. Position of the AND, DC, & ACSM: Nutrition and Athletic Performance. J Acad Nutr Diet. 2016 Mar;116(3):501528. [PubMed] Competitive athletes in a range of sports spanning the “Higher intakes may be strength-endurance indicated for short periods continuum during intensified training or when reducing energy intake.” “Higher protein intakes (2.3-3.1 g/kg/d) may be needed to maximize the retention of lean body mass in resistance-trained subjects during hypocaloric periods.” 1.2-2.0 g/kg - 16 - Caveats to accepting the position stands as gospel Although the position stands of the above organizations represent the weight of the evidence, this doesn’t mean that they are indisputable. The indicator amino acid oxidation (IAAO) technique is a validated method used for determining indispensable amino acids in humans.19 Recent studies using the IAAO technique have shown protein requirements greater than the low-end of the protein ranges listed in the position stands. Kato et al20 found that in endurance athletes on a training day had an estimated average requirement of 1.65 g/kg. More recently, Bandegan et al21 reported that the estimated average protein requirement in endurance-trained subjects in the 24-hour post-trained period was 2.1 g/kg. The latter findings call into question the recommendations of the current position stands on protein intakes for athletes, especially the allowance of intakes as low as 1.2-1.4 g/kg. Based on the current evidence, I would not recommend dipping below 1.6 g/kg for competitive athletes, or recreational athletes who take winning seriously. - 17 - Chapter 6: How much protein maximizes muscle gain? Let’s talk about growth for a moment Alright, so what about protein needs for muscle growth? This question is not all that simple. For maximizing muscle growth (also called muscle anabolism or hypertrophy), even the most carefully optimized protein intake is just part of the picture. In order to prime the physiological environment for growth, hypercaloric conditions (a caloric surplus) must be sustained. While muscle growth is indeed possible in caloric maintenance and deficit conditions, growth cannot be maximized unless a surplus of energy is consumed. Hypocaloric conditions compromise nutrient & energy availability. This suppresses anabolic signaling and muscle protein synthesis (MPS), ultimately compromising the rate of muscle growth. Hypocaloric conditions can tip the balance of turnover toward muscle protein breakdown (MPB). In contrast, hypercaloric conditions facilitate the opposite. Sustaining a caloric surplus drives muscle growth by not only increasing anabolic signaling and MPS, but also supporting the escalating demands of progressive resistance training volume. - 18 - Just what kind of caloric surplus is needed, you ask? The answer is, it depends on the population. Beginners and more advanced trainees have different requirements. The following table is a summary of the energy surplus guidelines from a recent paper I co-authored with Brad Schoenfeld:22 CALORIC SURPLUS GUIDELINE SUMMARY Population/training Magnitude of Nature of the surplus status the surplus Untrained/novice Approximately Greater potential benefit of a or deconditioned 20-40% above predominance of carbohydrate due to maintenance higher total energy surplus capacity. needs (~500Surplus should include a minimum 1000 kcal) protein dose of approximately 20-40 g (or at least ~0.4 g/kg of total bodyweight). Trained/more Approximately advanced; closer to 10-20% above maximum potential maintenance needs (~250-500 kcal) Lesser potential benefit of carbohydrate predominance due to lower total energy surplus capacity. Surplus should include a minimum protein dose of approximately 20-40 g (or at least ~0.4 g/kg of total bodyweight). Notes & caveats First and foremost, a caloric surplus for muscle gain must be built upon a foundation of sufficient total daily protein and energy intake. In general, for optimizing high-intensity fueling requirements of progressive resistance training, an energy surplus should focus on increasing carbohydrate. However, increased proportions of protein can be employed depending on how cautiously one wants to court the potential for concurrent fat gain. More advanced trainees closer to their potential have less room for surplus energy partitioning into lean tissue, and thus may choose to employ protein-focused surpluses. Regardless of training status, individuals cautiously avoiding fat gain might also benefit from this tactic. - 19 - Important side-note: a common misconception is that since resting muscle burns about 13 kcal/kg (6 kcal/lb) per day,23 only a tiny surplus is required to build muscle. Under that presumption, the surpluses in the above table might seem too large. However, aside from the research evidence showing otherwise,22 I would encourage you to head over to the July 2020 issue of AARR, and read the article titled, A pound of muscle burns [X] calories per day: facts, fallacies, & applications. In that article, I discuss the various components of energy expenditure increases involved with the process of building new muscle tissue. The resting value of 13 kcal/kg does not account for non-exercise & exercise activity increases, and thus should not be used for programming caloric surpluses for muscle growth. So, assuming we’ve got the right caloric surplus in place, maximizing muscle growth can be achieved with 1.6-2.2 g/kg (0.7-1.0 g/lb). This range is derived from Morton et al,24 who conducted the largest meta-analysis to date on the effect of protein supplementation on resistance training-induced gains in muscle mass and strength. I was fortunate enough to be one of the collaborators in this paper. Here’s a key passage from the discussion section which I snipped for brevity: “Here we provide significant insight by reporting an unadjusted plateau in RET-induced gains in FFM at 1.62 g protein/kg/day - 20 - (95% CI: 1.03 to 2.20). […] Given that the confidence interval of this estimate spanned from 1.03 to 2.20, it may be prudent to recommend ~2.2 g protein/kg/d for those seeking to maximise resistance training-induced gains in FFM.” With all this said, there are a few caveats to consider before taking the results as unassailable gospel. Note that this analysis excluded trials involving hypocaloric conditions, which have their own protein requirements (discussed in Chapter 8). It also did not focus specifically on highly trained, athletic, or competitive populations – let alone advanced trainees on ergogenic supplementation and/or drugs. Furthermore, protein needs based on total body mass are presumptive about body composition, when clearly there’s wide variability in the proportions of lean mass and fat mass between individuals. Nevertheless, Morton et al’s findings were echoed in subsequent research by Bandegan et al,25 who found a protein requirement of 1.7-2.2 g/kg in bodybuilders on a non-training day, using the indicator amino acid oxidation (IAAO) technique. Using the same method, Mazzulla et al26 reported that resistance-trained men required 2.01-2.38 g/kg. Current bodyweight vs. target bodyweight vs. lean mass Protein requirements based on total bodyweight predominate the peer reviewed literature; it’s actually quite rare to find - 21 - publications that issue recommendations based on lean mass (typically denoted as fat-free mass or FFM). Total bodyweightbased protein recommendations are typically issued under the presumption of normal-weight individuals. The pitfall here is that it’s possible to overshoot or undershoot estimated needs if someone is highly over- or underweight. A simple solution to this, which also circumvents the need to estimate body composition, is to base protein intake on goal bodyweight or target bodyweight. Use your current weight to estimate protein needs if you’re seeking to maintain current weight. Otherwise, base your estimations on target bodyweight. This is an effective way of approximating lean mass, with a built-in margin of safety. A quick-and-dirty estimate of protein needs that doesn’t factor bodyweight into the calculation is basing your protein gram target on your height in centimeters (multiply inches by 2.54). If you run the numbers on this, you’ll find that it’s in the right ballpark for taller folks, but shoots high for shorter individuals. As such, it tends to overestimate the protein needs of women who are not particularly tall. For those who are hell-bent on basing protein intake on lean mass or fat-free mass (FFM), a reasonable range for maximizing muscle growth is 1.8-2.6 g/kg FFM. This range is - 22 - derived from IAAO data in recent studies I discussed in the October 2019 issue of AARR. To simplify things, a target of 2.2 g/kg FFM (1.0 g/lb FFM) shoots right in the middle of the ‘optimal’ range. This is a safe baseline target for the goal of muscle gain. Like any programming variable, this would need to be put to trial and adjusted as needed. Summing up • Maximizing muscle growth requires a sustained caloric surplus (muscle growth can still occur without a surplus, it just won’t be maximized). • Based on the collective longitudinal research that directly assessed body composition, total daily protein intake for maximizing muscle gain is 1.6-2.2 g/kg of total bodyweight (0.7-1.0 g/lb).24 • Women typically carry significantly more body fat than men, so shooting lower on this range might be more appropriate as a starting point from which to adjust according to results. • Remember to use target bodyweight if you’re highly overor underweight. • Alternatively, 2.2 g/kg FFM (1.0 g/lb FFM) can be put to trial and adjusted according to individual response. - 23 - Chapter 7: How should protein be distributed through the day to maximize muscle gain? For gaining muscle, distribution (pattern, frequency, or spread) of protein doses through the course of the day appears to matter, especially if the goal is to maximize muscle growth. To the best of our knowledge, pushing maximal growth is best accomplished by having protein in similar doses through the course of the day,27 from waking to pre-bed.28 Each dose should maximally stimulate muscle protein synthesis (MPS). Muscle protein breakdown (MPB) is the other side of protein turnover. Maxing-out MPS (as opposed to minimizing MPB) is the more important target since changes in MPS are 4-5 times greater than MPB in response to exercise and feeding..29 Both processes occur in constant, dynamic cycles. Muscle growth occurs as a result of MPS exceeding MPB over time. Recent research has refuted the longstanding presumption that 20-25 g high-quality protein (containing >2 g leucine) elicits a maximal anabolic response. Macnaughton et al30 reported greater MPS with 40 vs 20 g whey after a high-volume resistance training session, and Park et al31 reported greater MPS with 70 vs 35 g beef protein within a mixed macronutrient meal. Collectively, the current evidence suggests a minimum of - 24 - 4 feedings dosed at approximately 0.4-0.55 g/kg per meal (the full text detailing the rationale of this distribution is available here).32 Now, is it possible that muscle growth can be maximized with a lower number of feedings than 4 per day, or a more skewed/less even distribution of meals? Yes, it’s possible. That’s one of the gray areas of research with room for more investigation. I would even speculate that 3 protein feedings per day versus 4 (or more) is not likely to be meaningful outside of competitive conditions where very small differences can determine different placings. However, I would also speculate that less than 3 protein feedings per day is not likely to maximize hypertrophy from a practical standpoint (gastrointestinal issues associated with huge meals), as well as a mechanistic standpoint (the anabolic “ceiling” seen in the limited MPS elevations per feeding). I’ll reiterate that muscle growth is possible with low feeding frequencies, including purposely narrowed feeding windows. However, merely achieving some degree of muscle growth is a distinctly different goal from maximizing rates of muscle growth. Protein feeding frequency & distribution will remain an area of controversy as long as there’s a lack of direct comparisons of different meal frequencies in resistance trainees in trials that measure body composition changes over - 25 - time. In the meantime, the following table provides total and per-meal dosing specifics. PROTEIN DISTRIBUTION FOR MAXIMIZING MUSCLE GROWTH Bodyweight Total Daily Protein Dose Per Meal (3 Meals) Dose Per Meal (4 Meals) 50 kg (110 lb) 80-110 g 27-37 g 20-27 g 55 kg (121 lb) 88-121 g 29-40 g 22-30 g 60 kg (132 lb) 96-132 g 32-44 g 24-33 g 65 kg (143 lb) 104-143 g 35-48 g 26-36 g 70 kg (154 lb) 112-154 g 37-51 g 28-38 g 75 kg (165 lb) 120-165 g 40-55 g 30-41 g 80 kg (176 lb) 128-176 g 43-59 g 32-44 g 85 kg (187 lb) 136-187 g 45-62 g 34-47 g 90 kg (198 lb) 144-198 g 48-66 g 36-50 g 95 kg (209 lb) 152-209 g 51-70 g 38-52 g 100 kg (220 lb) 160-220 g 53-73 g 40-55 g 110 kg (242 lb) 176-242 g 59-81 g 44-60 g 120 kg (264 lb) 192-264 g 64-88 g 48-66 g 130 kg (286 lb) 208-286 g 69-95 g 52-71 g - 26 - Chapter 8: How much protein maximally preserves muscle while losing fat? Generally speaking Protein requirements increase depending upon the severity of the caloric deficit, and the degree of leanness of the dieter.33 In hypocaloric (dieting) conditions, untrained individuals with ample/excessive body fat lose a greater proportion of their bodyweight from fat mass compared to lean, trained individuals, who stand to lose more lean mass. In sustained eucaloric (weight maintenance) conditions,34 and even in hypercaloric (weight gain) conditions,35 low protein intakes still can jeopardize muscle mass. Athletes’ requirements The threat to muscle preservation in caloric maintenance and surplus conditions via inadequate protein intake is amplified in hypocaloric conditions. Higher protein intakes combined with resistance training is the antidote to muscle loss in hypocaloric conditions. A recent (2019) review by Hector and Phillips.36 concluded that an appropriate range of protein intake for athletes in hypocaloric conditions is 1.6-2.4 g/kg. - 27 - Pushing the envelope Helms et al37 conducted a systematic review that concluded that 2.3-3.1 g/kg of fat-free mass (FFM) was appropriate for resistance-trained subjects in hypocaloric conditions, with needs escalating according to the severity of the deficit and leanness level. However, out of the six studies in the review, only three examined highly-trained competitive athletes, and only one study examined competitive bodybuilders. The latter study on bodybuilders by Mäestu et al38 reported that the precontest protein intake of drug-free, world-class bodybuilders ranged 2.48-2.68 g/kg. Chappell et al13 reported that elite-level drug-free bodybuilding men & women who placed in the top-5 had pre-contest protein intakes of 3.3 & 2.8 g/kg, respectively. While these data are observational and thus are incapable of establishing causation, they’re still valuable. As the late Jim Rohn wisely said, “Success leaves clues.” Summing up Boiling down protein requirements to maximize muscle retention while dieting, it might help to delineate 3 population tiers, thus 3 sets of recommendations: • For athletic populations in general, the collective literature shows that 1.6-2.4 g/kg (0.72-1.09 g/lb) of total bodyweight - 28 - is appropriate.24,36 You can base this calculation on target bodyweight for those who are highly over- or underweight. • For relatively lean, resistance-trained competitive athletes, an appropriate range is 2.3-3.1 g/kg (1.04-1.41 g/lb) of fatfree mass. Based on a mean body fat level of roughly 18% among the subjects in the treatment groups of Helms et al’s systematic review (which included men and women),37 this translates to 1.9-2.5 g/kg (0.86-1.14 g/lb) of total bodyweight. • For those pushing the envelope of leanness in the pre-contest phase of physique competition, 2.4-3.3 g/kg (1.1-1.5 g/lb) of total bodyweight appears to optimize this goal at the elite levels.12,13 - 29 - Chapter 9: How should protein be distributed through the day to maximize muscle retention while dieting? It’s logical that the protein distribution pattern that maximizes muscle growth under ideal (surplus/hypercaloric) conditions is likely to be the same pattern that maximizes muscle retention under compromised (deficit/hypocaloric) conditions. However, there’s evidence that the goal of muscle retention has a greater allowance for suboptimal/skewed protein intake patterns, which we’ll cover in a moment. Mind the hierarchy First things first – keep in mind that in the hierarchy of importance, total daily protein intake strongly holds the top spot. As covered in the previous section, for physically active and athletic individuals, which is most of you reading this, 1.62.4 g/kg (0.72-1.09 g/lb) of total bodyweight is an appropriate target for total daily protein. 24,36 This is nearly identical to the range that maximizes muscle growth (1.6-2.2 g/kg).24 The distribution pattern likely to maximize muscle growth in ideal conditions is a minimum of 4 feedings dosed at approximately 0.4-0.55 g/kg per meal.32 The latter scheme, including a - 30 - relatively even spread of protein doses from waking to prebed28 probably holds true for maximizing muscle retention. Asymmetric control Back to the first point. There appears to be an asymmetric control system regarding muscle growth versus muscle retention. Growth – especially beyond “newbie gains” – is more difficult to accomplish than merely preserving existing muscle mass. The mechanisms underlying this observation are poorly understood. Perhaps it’s simply more energetically expensive to fuel the molecular processes involved with muscle growth. The body’s homeostatic drive resists changes; it fights to neutralize shifts in the status quo, perceiving continued muscle growth beyond normal/baseline levels as a threat to survival, for reasons not definitively known. Compounding this, there’s the messy matter of striking a balance (or even a compromise) between maximizing the rate of muscle growth while minimizing excessive gains in body fat. Regardless of the above speculations, the big point to take home is that muscle retention in hypocaloric conditions has been observed in the absence of an idealized protein distribution through the day. This has been most saliently - 31 - demonstrated in recent time-restricted feeding (TRF) studies involving a formal resistance training protocol, where protein intake was adequate (at least 1.6 g/kg) and equated between the groups in the comparison. Emerging data To date, there are three studies that fit the above criteria. An 8week study by Moro et al39 was dubbed the Leangains study since it tested the 16:8 fast:feed cycle popularized by Martin Berkhan. A 3-meal TRF pattern consisting of 16-hour fasting/8hour feeding cycles (16/8 TRF, meals occurred at 1 pm, 4 pm, and 8 pm) was compared with a conventional/control diet with 3 meals distributed within 12-hours (meals occurred at 8 am, 1 pm, and 8 pm) in resistance-trained men. Training occurred between 4 pm & 6 pm. No significant changes in lean mass occurred in either group. Interestingly, greater fat loss occurred in TRF. A subsequent study by Tinsley et al40 conducted a similar comparison in resistance-trained women. Meals in TRF occurred between 12 pm and 8 pm, while the control group self-selected their meal times throughout the day without timeframe restrictions. Training occurred between 12 pm & 4 - 32 - pm. Lean mass increased in all groups, but fat loss only occurred in the TRF. With both of these studies being freeliving designs rather than tightly controlled in-patient designs where all intake is lab-provided and subjects are closely supervised, the most plausible explanation was that the superior fat loss of TRF was due to lower energy intake. The latter speculation is supported by existing research showing a 20-30% energy intake decrease in ad libitum (unrestricted) TRF conditions.41 Most recently, Stratton et al42 reported a lack of difference between the effects of an 8-hour TRF and a normal diet in recreationally trained subjects undergoing resistance training at a 25% caloric deficit for 4 weeks. Body composition changes (lean mass preservation and fat loss) and strength/power improvements were similar in both groups. This lack of fat loss advantage breaks the momentum of the previous TRF + resistance training studies.39,40 It’s notable that Stratton et al’s study involved more rigorous compliance measures than in previous research. Subjects were interviewed during each workout, where dietary logs were reviewed at the beginning of each week. Furthermore, a 4-compartment model that employed dual X-ray absorptiometry (DXA), bioelectrical impedance analysis (BIA), air displacement plethysmography - 33 - (ADP, also called BOD POD), and ultrasonography was used to assess body composition. Summing up Based on the limited research thus far, we can conclude that an 8-hour feeding window is apparently not a threat to muscle retention. A welcome side-effect is that TRF has the potential to facilitate fat loss – but not likely via any “magic” aside from less opportunity to eat. A fat loss advantage of TRF + resistance training has been seen in two39,40 out of the three studies thus far examining the question. 39,40,42 So, the potential advantage is there, but it’s not without dispute. With that said, there are unanswered questions worth pondering. Can 2-3 protein-rich meals per day be as effective as 4 or more protein-rich meals for the goal of retaining muscle mass in hypocaloric/dieting conditions? I’m confident in saying that 3 meals within an 8-hour period can do the trick, since we have consistent evidence of that in resistance trainees.39,40,42 Furthermore, a meta-analysis by my colleagues and I found that in hypocaloric conditions, meal frequency overall had minimal influence on body composition even in a general lack of resistance training across studies included in the metaanalysis.43 - 34 - At this point, we can only speculate what the limits of suboptimality are in terms of protein distribution through the day. When forming guidelines for application, practicality and tolerability diminishes sharply below 3 meals per day for most individuals. Nevertheless, the question is interesting, and also relevant to dieters with a preference for lower meal frequency. Based on my observations, I’d speculate that as low as 1-2 protein-rich meals per day – although not feasible for many – can still retain muscle mass, given that resistance training and macronutrition are sound, and the energy deficit is not severe (not allowing more than an average of 1% of BW to be lost per week). - 35 - Chapter 10: How does protein intake influence recomposition? Recomposition, nicknamed “recomp” by the fitness community, is the coveted holy grail. It’s important to keep in mind that recomp capacity diminishes alongside the progression of an individual’s training status.44 Put more simply, recomp happens more dramatically in untrained (or previously trained) folks with excess body fat. The closer you are to your potential, the less available margin there is for recomp. With advancing proximity to your potential, recomp becomes an impractical target. Advanced trainees and most later-intermediates are best served by focusing on one goal at a time: fat loss or muscle gain, not both simultaneously. For an in-depth discussion of what distinguishes beginners, intermediates, and advanced trainees, refer to the August 2019 issue of AARR. The hierarchy of recomp capacity, from greatest to least, is: 1) formerly trained folks with excess body fat, 2) overweight novices, 3) intermediates, 4) advanced trainees. I’d like to briefly discuss the recomp advantages of formerly fit folks. This is the profile of those who make the most dramatic before-after - 36 - transformations, especially if they over-ate while remaining sedentary for a prolonged period. There are at least two potential mechanisms underlying this ability of the formerly fit to dramatically rebound back into shape. Training increases the number of myonuclei within the muscle fibers.45 Strength & size gains can thereby be expedited via increased mitochondrial remodeling. Training-induced myonuclear increases are resistant to apoptosis (cell death), and remain intact despite extended time off from training.46 Another potential mechanism is the persistence of traininginduced increases in capillarization (an increase in vascular networks within the muscle).47 As a result, the rebound gains in previously trained individuals are significantly faster than the gains seen in beginners who’ve never trained before. When you combine these mechanisms with greater neural efficiency reducing the learning curve of the exercises, it’s clear that the recomp advantages of previously trained folks (who have gained body fat) put them at the top of the hierarchy. Summing up Recomp does in fact occur across a range of populations. However, recomp capacity diminishes alongside increased - 37 - training status. Another consistent finding is that higher protein intakes are more conducive to recomp. A nicely done review paper by Bakarat et al44 is the first of its kind in the peer reviewed literature to specifically examine the recomp phenomenon. Here are the authors’ practical application points, relayed verbatim from the manuscript (proteinoriented points italicized by me): • Implement a progressive RT regimen with a minimum of 3 sessions per week. • Tracking rate of progress, and paying attention to performance and recovery can be important tools to appropriately adjust training over time. • Consuming 2.6-3.5 g/kg of FFM (fat-free mass) may increase the likelihood or magnitude of recomposition. • Protein supplements (i.e., whey and casein) may be used as a means to increase daily dietary protein intake as well as a tool to maximize muscle protein synthesis. This may be of greater importance postworkout as a means to maximize the recomposition effect. • Prioritizing sleep quality and quantity may be an additional variable that can significantly impact changes in performance, recovery, and body composition. - 38 - Chapter 11: How does protein timing (pre, during, and post-exercise) impact athletic performance? I’ll begin by saying that protein per se (especially compared to total caloric intake and carbohydrate intake in particular) has minimal utility as a fuel for enhancing athletic performance. Although we can end this discussion here, it’s useful to know the key pieces of research on this topic. It’s also useful to know the primary rationale, which is to prevent muscle protein breakdown (MPB), which can indirectly impact performance if it manifests as a loss of muscle mass. Another justification for the intake of protein or amino acids during exercise is to prevent central fatigue, but this is a hypothesis that has largely failed across multiple studies.48,49 Pre-exercise On the pre-exercise front, the only study to date which directly compared protein with other macronutrients on endurance performance was back in 2002 by Rowlands and Hopkins.50 Twelve competitive cyclists ingested either a high-fat, highcarbohydrate, or high-protein meal 90 minutes prior to a test involving sprinting and 50 km performance. No significant performance differences were seen between the treatments - 39 - compared. As for strength performance, to my knowledge, there is no study specifically comparing pre-exercise protein with other macronutrients. During exercise A crucial yet commonly overlooked factor influencing the ergogenic potential of intraworkout nutrition (protein or other) is the presence versus absence of preworkout nutrition, which from a bioavailability standpoint, can function as intraworkout nutrition, considering the time course of digestion and absorption, which can last several hours. In any case, in the most recent meta-analysis to examine the effect of protein & carbohydrate co-ingestion versus carbohydrate-only on endurance performance, Nielsen et al49 reported that when carbohydrate intake was equated, the condition with more protein increased endurance capacity. However, this was due to greater total energy intake. In comparisons that equated total calories, there was no endurance performance advantage of a carb-protein mix versus carbs alone.50 Again, depending on how close to the start of the bout, preworkout nutrition can function as intraworkout nutrition. For example, when 45 grams of whey is ingested alone, it takes about 45 minutes for circulating amino acid levels to peak, and another 2 hours to return to baseline levels.51 - 40 - As such, the question of intraworkout protein intake only applies to fasted resistance training, which has limited relevance to the real world. Given this far-fetched scenario, a case can be made for intraworkout nutrition, and indeed there is evidence of its benefit for resistance trainees. Bird et al52 found that a beverage containing carbohydrate and essential amino acids suppressed MPB. This finding is potentially relevant to those with no option except to train immediately upon waking, where strength gains would be compromised with continued net losses in muscle protein. I’ll reiterate that this was observed in fasted conditions minus any pre-exercise nutrition, which is far-fetched for resistance training. For those interested in a deeper dive into the research on intra-workout protein consumption & endurance performance, it’s been thoroughly examined in the July 2020 issue of AARR. Post-exercise A recent meta-analysis by Craven et al53 reported that protein co-ingested with carbohydrate does not expedite post-exercise glycogen resynthesis (or subsequent exercise performance) compared to carbohydrate alone, especially when the dosing is at or near 1.0 g/kg/hr. This echoes the recommendations of recent position stands on post-exercise carbohydrate dosing (1-1.2 g/kg/hr) for maximizing rates of glycogen replenish- 41 - ment under time-constrained conditions where competitive endurance bouts are separated by less than 8 hours.18, 54 Summing up In perhaps the most anticlimactic sum-up ever, protein timing is a relative non-factor in relation to athletic performance. It can even be said that maximizing athletic performance is partially a matter of not letting your protein intake get in the way of your carb intake. Although there’s the potential benefit of intraworkout protein intake for the purpose of mitigating muscle protein breakdown to indirectly preserve strength, this only applies to an absence of preworkout (and/or postworkout) nutrition. The “anabolic window” concept will be addressed in the next section where protein timing & muscle growth are discussed. - 42 - Chapter 12: How does protein timing (pre, during, and post-exercise) impact body composition? Let’s first establish the reality that total daily nutrient & calorie intake have the greatest impact on body composition. In the ISSN’s position stand on diets & body composition,33 I called the sum of those totals “the cake,” while timing of its constituents is “the icing.” Protein timing is of distant secondary importance. It’s common for folks to fall prey to marketing hype and guruism that puts timing up on a magic pedestal, in opposite order of importance. Get the cake right first, then you can apply the icing. Pre-exercise There’s an interesting line of research showing that protein taken prior to exercise results in greater increases in energy post-exercise expenditure compared to carbohydrate.55,56 However, short-term differences in energy expenditure don’t guarantee advantages in body composition change over the longer-term. A direct comparison of pre- versus post-exercise protein intake by my colleagues and I failed to show any - 43 - meaningful body composition advantage to either protocol.57 I’ll touch upon this one again in a moment. During exercise As mentioned in the previous section, Bird et al52 showed that lifting-induced muscle protein breakdown (MPB) was suppressed by a liquid carb/essential amino acid mixture. The same caveat applies: the threat of MPB is likely limited to fasted training sessions. Even so, the body is often smarter than we give it credit for. Deldicque et al58 reported a greater anabolic signaling response to the post-exercise meal after fasted exercise, compared to exercise in fed conditions. It’s possible that despite the inherently catabolic nature of training in fasted conditions, the body amplifies its use of the post-exercise feeding, to supercompensate, and preserve homeostasis after perceiving an energy crisis (thus a threat to survival). Nevertheless, the schedule constraints of some individuals might necessitate immediate commencement of the training session upon waking. In this case, intraworkout protein might be justified if the goal is to pull all theoretical stops to maximally protecting against muscle catabolism. The provision of default hydration can’t hurt either, given the obvious refrain from drinking fluids during sleep. - 44 - Post-exercise Again, we have to think in terms of cake versus icing. Regarding body composition effects, we’re definitely in icing territory. The “post-exercise anabolic window” concept was brought to the general fitness & bodybuilding audience in the early 2000’s, thanks to Ivy and Portman’s research, and their popular paperback Nutrient Timing: The Future of Sports Nutrition.59 The general premise was that the timing of specific nutrients immediately post-exercise (quickly absorbed protein and carbs) could make or break muscular gains. The implication was that nutrient timing within the “anabolic window” (within an hour postexercise) was more important than totals by the end of the day.60 This principle got so deeply ingrained in the mantra of trainees and coaches, it became accepted as a matter of fact. However, Ivy & Portman’s premises were based on short-term anabolic response studies that measured muscle protein synthesis (MPS) and/or glycogen resynthesis. Longitudinal studies (lasting several weeks or months) examining protein timing’s effects on muscular adaptations to resistance training told a different story. Several studies published over the decade following Ivy & Portman’s book yielded highly equivocal results, casting - 45 - considerable doubt upon the postexercise anabolic window concept. In light of these mixed results across multiple protein timing studies, my colleagues and I conducted a meta-analysis of the relevant research.61 We defined protein-timed conditions as protein ingestion within an hour of either side of the resistance training bout, while non-timed conditions involved a minimum of 2 hours of protein neglect on both sides of the training bout. Basic analysis (not accounting for covariates such as total daily protein) showed a minor advantage of protein timing on hypertrophy. However, regression analysis found that this effect was due to greater total daily protein intake in proteintimed conditions compared to the non-timed conditions (1.66 vs. 1.33 g/kg, on average). A subanalysis of the studies that did equate total daily protein still failed to show an advantage of protein timing. So, we concluded that if an anabolic window for protein intake exists, it appears to be greater than the one-hour before and after a resistance training session. Summing up Keep in mind that the following bullet points sum up protein’s role in this particular topic (timing for athletic performance). - 46 - It’s not a comprehensive treatise on nutrient timing; this book is focused on protein. The role of carbohydrate timing in exercise performance is covered in-depth in the April & May 2020 issues of AARR. • Pre-exercise protein has the potential to increase postexercise energy expenditure to a greater degree than carbohydrate. Whether these short-term effects can manifest as body composition improvements is speculative in the absence of data. • Protein and carbs consumed during exercise (intraworkout) can suppress muscle protein breakdown, but the relevance and applicability of this is limited to overnight-fasted conditions, and the absence of preworkout nutrition. • Intraworkout nutrient consumption is not necessarily beneficial in the context of a preworkout meal already being digested and absorbed (in which case it can actually be redundant & superfluous). If intraworkout intake is personally preferred, that’s fine. It generally won’t hurt, and can help in certain cases. • Instead of focusing on a narrow post-exercise “anabolic window,” the period between the pre- & postworkout meal (the periworkout period) should be the focus.62 - 47 - • The anabolic effect of a protein-rich meal is 3-5 hours, potentially longer, depending on meal size and composition. The preworkout meal in many cases can dictate the postworkout environment. • Determined by preference & tolerance, liquid or solid meals can be consumed at any point within the 3-5 hour periworkout period. • Protein dosing in both the pre- & postworkout meals should maximize the anabolic response (0.4-0.55 g/kg).32 • Note the periworkout period in green, which varies in length according to individual variables such as meal size, form, and composition: PERIWORKOUT TIMING MODEL: THE “ANABOLIC GARAGE DOOR OF PEACE” - 48 - Chapter 13: How does BCAA supplementation affect body composition? Well hyped, well loved Branched-chain amino acids (BCAAs) are one of the most popular supplements among recreational and competitive athletes. Chappelle et al63 recently reported that among highlevel, drug-free competitive bodybuilders, BCAAs were the third most commonly used supplements, with the first and second being protein powder and multivitamins, respectively. Mechanistic rationale The BCAAs are leucine, isoleucine, and valine. The BCAAs happen to be 3 of the 9 essential amino acids (EAAs). Of the BCAAs, Leucine is considered to play a key role in driving muscle protein synthesis (MPS), suppressing muscle protein breakdown (MPB), and activating anabolic signaling.64 The “leucine threshold” where MPS is stimulated above resting levels, is a loosely defined dosing threshold of at least 1-2 g leucine, with the upper end being more applicable to older subjects.65 Jackman et al66 showed that post-exercise BCAA supplementation, even in the absence of other amino acids, increases myofibrillar MPS and mTORC1 signaling. BCAA - 49 - supplementation has also been shown to reduce delayed onset muscle soreness (DOMS),67 which implicates BCAA in the suppression of exercise-induced muscle damage. But does it work? Short-term anabolic response data is useful for generating hypotheses and investigating potential mechanisms. However, what matters most is whether BCAA supplementation actually enhances muscle growth or fat loss under relevant conditions. This can only be investigated via longitudinal studies lasting several weeks or months. To date, Dudgeon et al68 conducted the only study to show superior body composition effects of BCAA supplementation versus control conditions in resistance trainees consuming optimal total daily protein (≥1.6 g/kg). However, this study was so full of errors and inconsistencies, my colleagues and I were compelled to point them out in a formal letter.69 In contrast to Dudgeon et al’s findings,68 there are at least four studies involving a structured resistance training protocol that found no significant effects of BCAA supplementation on body composition or strength compared to placebo.70-74 A recent study 16-week study by Ooi et al75 did not involve resistance training, but compared 3 groups in a 500 kcal deficit: - 50 - 1) standard-protein (14%) plus placebo as a control condition, 2) standard protein plus BCAA dosed at 0.1 g/kg/day, and 3) high-protein (27%) plus placebo (HP). Fat mass loss was similar across the groups. Although the differences in lean mass loss didn’t reach statistical significance, the high-protein group lost the least lean mass, with the BCAA and control groups coming in second and third in this regard. These results align with the concept that the whole (intact protein) is more than the sum of – some of – its parts (BCAAs). Even the non-essential amino acids (NEAAs) can function integrally with the other components of the protein. To illustrate this, I’ll quote an excerpt from a thought-provoking review by Hou et al:76 “Although EAA and NEAA had been described for over a century, there are no compelling data to substantiate the assumption that NEAA are synthesized sufficiently in animals and humans to meet the needs for maximal growth and optimal health. NEAA play important roles in regulating gene expression, cell signaling pathways, digestion and absorption of dietary nutrients, DNA and protein synthesis, proteolysis, metabolism of glucose and lipids, endocrine status, men and women fertility, acid–base - 51 - balance, antioxidative responses, detoxification of xenobiotics and endogenous metabolites, neurotransmission, and immunity.” Summing up So, does BCAA supplementation enhance body composition? That’s highly unlikely – especially given sufficient total daily protein intake. With scant exception, studies involving resistance training fail to support the BCAA supplementation’s effectiveness for either body composition or strength improvements beyond placebo. Evidence aside, from a logical and theoretical standpoint, BCAA supplementation is difficult to justify. High-quality dietary protein sources are already composed of 18-26% BCAA. To quote Kevin Finn, “Taking BCAAs is like turning the sprinklers on when it’s raining out.” A better alternative to taking BCAA is to spend your money on high-quality protein. You’ll get the BCAAs, the rest of the EAAs, and multiple nutritive cofactors within the food matrix that can enhance muscle growth, health, and exercise performance. It’s a win-win: you save money and get more benefits. - 52 - Chapter 14: How does a high protein intake impact ketosis? Under normal, non-dieting conditions, circulating ketone levels are low (<3 mmol/l). Aside from completely fasting, ketosis is attained by restricting carbohydrate to a maximum of ~50 g or ~10% of total energy, with the predominance of energy intake from fat (~60-80% or more, depending on degree protein and carbohydrate displacement).33 This brings circulating ketone levels to a range of ~0.5-3 mmol/l. The primary ketone in the blood is β-hydroxybutyrate (BHB). Ketogenic diet proponents claim rather wishfully that greater degrees of ketosis provide greater benefits for dieters. In their book The Art and Science of Low Carbohydrate Performance,77 keto research pioneers Jeff Volek and Stephen Phinney recommend blood BHB levels ranging 0.5-3.0 mmol/l to achieve “optimal fuel flow during keto-adaptation.” They also specifically define “nutritional ketosis” as a BHB range of 1.0-3.0 mmol/l. They also recommended a relatively low protein intake (0.6-1.0 g kg of lean mass). Readers took this as gospel, and avoided high protein intakes for fear of proteinmediated gluconeogenesis kicking them out of ketosis. - 53 - Moderate protein in keto diets has been espoused in the peer reviewed literature as well. Quoting Paoli et al: 78 “…it is not correct to equate a ketogenic diet with a high protein diet, because the state of the art ketogenic diets are normoproteic thus the daily amount of protein is about 1.2-1.5 g of protein per kg of body weight.” However, there are enough data from controlled intervention trials to confidently say that protein intakes beyond traditional recommendations are still permissive of BHB levels that would make keto proponents happy. Summing up Here are the data of studies demonstrating that high protein intakes can still allow blood ketone levels that fall within the commonly cited range that qualifies as ketosis (0.5-3.0 mmol/l).79-81 Note that two of the three studies below yielded the magical range of blood ketones (1.0-3.0 mmol/l) promoted by keto connoisseurs. Please note that I’m being completely facetious when I say magical. Whether ketogenic dieting is good, bad, or neutral depends on individual goals and preferences. - 54 - Although it’s slightly off-topic from this book, it’s worth mentioning that there’s nothing inherently superior about ketogenic dieting vs. non-ketogenic dieting for altering body composition or improving performance. See the keto FAQ in the August 2018 issue of AARR, or the ISSN Position Stand on Diets & Body Composition for more details on ketogenic diets. STUDIES SHOWING KETOSIS DESPITE HIGH-PROTEIN DIETS Publication Population Protein Carb Blood intake intake ketone level Wilson JM, et al. 25 college-aged 1.7 g/kg 31 g/day 1.0 mmol/l J Strength Cond resistanceRes. 2020 trained men Dec;34(12):3463374. [PubMed] Burke LM, et al. J 21 elite race Physiol. 2017 May walkers 1;595(9):27852807. [PubMed] Volek JS, et al. Metabolism. 2016 Mar;65(3):100-10. [PubMed] 2.2 g/kg 33 g/day 1.8 mmol/l 20 elite ultra2.1 g/kg marathoners & ironman distance triathletes 82 g/day 0.7 mmol/l - 55 - Chapter 15: How does protein restriction influence longevity? Many people have heard the claim that protein restriction increases lifespan. Many have accepted it unquestioningly, and filed it into their belief system. It’s a convenient, common claim for folks whose dietary ideology makes getting optimal protein targets somewhat of a pain in the butt. After all, optimizing protein intake can require extra effort, strategy, and expense. It would be much simpler (and cheaper) to just skate by on a low protein intake, thinking you’re getting the added benefit of greater longevity. Rogue offspring of caloric restriction Protein restriction is an offshoot of caloric restriction (CR), which is a prolifically studied tool for promoting longevity in animal models. However, a stiff challenge to the protein restriction for longevity model is that it’s not rooted in human research. For example, glucose and amino acid restriction have been shown to extend the lifespan of Saccharomyces cerevisiae, a single-celled fungus, better known as brewer’s yeast.82 An abundance of longevity experiments have been done on yeast, due to its short lifespan and ease of trial repeatability. - 56 - Flies, worms, and rodents (oh my) Lifespan increases from amino acid or protein restriction have also been seen in flies, worms, and rodents. In non-human primates, however, the findings have been equivocal, and protein restriction cannot necessarily be separated from calorie restriction (CR). Notably, in rhesus monkeys, the Wisconsin National Primate Research Center showed significantly lower mortality in the CR group compared to controls, whereas the National Institute on Aging reported no significant difference.83 Keep in mind that animal data do not automatically apply to humans. Animal data are, at best, hypothesis-generating food for thought, subject to replication (or refutation) by controlled intervention trials on…you guessed it, humans. A tale of two monkey studies This leads us to human research on protein restriction. Like the non-human primate data, human data are also equivocal. Exemplary of this mixed bag results is Levine et al’s analysis84 of the Third National Health and Nutrition Examination Survey (NHANES III) data combined with follow-up data from the National Death Index. Moderate protein intakes (10-19% of calories from protein) and high protein intakes (≥20% of - 57 - calories from protein) had higher risks of diabetes-related mortality than the low protein intakes (<10% of calories from protein) group. Among those aged 50-65 years, higher protein intakes increased risks of all-cause and cancer-related mortality. However, in subjects aged 66 years and older, the high protein intakes were associated with the opposite effect – lower all-cause and cancer-related mortality. In addition to the dichotomous results, it’s worth reiterating that Levine et al84 reported observational data, incapable of demonstrating cause-and-effect. The authors conceded that a major limitation with their findings is that they were based on a single 24-hour dietary recall, followed up by as much as 18 years of mortality assessment. Beloved BCAA escapes the chopping block Branched chain amino acid (BCAA) intake has been implicated in higher protein intakes’ purported antagonism of health and longevity via driving the mTOR pathway, and somehow facilitating insulin resistance.85 Again, this concern is based on rodent data and observational leaps. It’s poorly justified, given the importance of these metabolic pathways in preventing sarcopenia. In contrast to the observational data drawing an - 58 - association between circulating BCAA levels insulin resistance in obese and diabetic patients,85 human intervention data failed to support this. In a recent crossover study on obese, prediabetic subjects, Woo et al86 found that high-dose BCAA supplementation (20 g/day) did not affect glucose metabolism. Muscle-centric approach as an antidote to aging A recent review by Burd et al87 presented a muscle-centric perspective of optimizing protein intake through the lifespan for the purpose of optimizing health. The RDA (0.8 g/kg) fails to optimize the retention of muscle mass, especially in the elderly. The range cited by Burd et al (1.2-1.5 g/kg)87 is very close to the 1.2-1.6 g/kg reported by Phillips et al7 for preserving muscle mass and optimizing metabolic health through the aging process in the general population. Importantly, Burd et al87 and Phillips et al7 based their conclusions on randomized controlled trials that consistently favor higher protein intakes for improving body composition and a wide range of cardiometabolic health parameters. This is in stark contrast to the animal data (much of it from rodents and invertebrates) and observational data (incapable of showing causation) suggesting the low-balling of protein intake for longevity. - 59 - Sarcopenia (age-related progressive loss of muscle mass and function) is one of the greatest health threats faced by older adults.88 A loss of muscle mass generally begins at ~50 years and progresses at a rate of ~0.8% per year.87 Strength decline is ~2–3% per year. At 70 years of age, there is the potential for a 16% loss of muscle mass and 50% drop in strength compared to younger adulthood. The good news is that sarcopenia is preventable and treatable with nutrition and training interventions. A proactive, musclecentric approach needs to replace the unfortunately common reactive approach, where advancing age finally catches up with a sub-optimal protein intake. If anything, age-related anabolic resistance (a decreased anabolic response to protein feedings) actually warrants an increase in protein intake with advancing age, not a decrease.7 Summing up: I’ll end off by quoting an excerpt from the December 2017 issue of AARR, which I feel sums up this topic well: “For increasing longevity, the case for protein and amino acid restriction is relatively weak, disjointed, and hypothetical - 60 - compared to the case for raising protein intake above the currently recommended amounts by public health organizations. The benefits of protein restriction are rooted in animal data, whose results are largely divergent from controlled human data examining a broad spectrum of endpoints that define healthy aging. Given the interplay of factors involved with the development of the age-related pathologies, it’s not surprising that the bestsupported defense against sarcopenia and its related conditions is a two-pronged intervention: optimized dietary protein intake and exercise (especially resistance exercise).” - 61 - Chapter 16: How does advanced age impact protein utilization and dosing requirements? Anabolic resistance is characterized by a diminished muscle protein-synthetic (MPS) response to protein/amino acid feeding & exercise.89 This is typically an age-related phenomenon that contributes to the decline of muscle mass and function. Precise mechanisms are yet to be definitively elucidated, but the “use it or lose it” principle (applied to muscle), is a prime contributor. Decreased protein and total energy intake, in addition to increased physical activity are the main interplaying culprits in the development of anabolic resistance. Per the hierarchy of importance regarding protein variables, total daily intake is at the top. According to expert panel positions (PROT-AGE and ESPEN),6 individuals over age 65 have the following protein requirements: 1.0-1.2 g/kg for healthy folks, 1.2-1.5 g/kg in cases of acute or chronic illnesses, and 2.0 g/kg for those with severe illnesses, injuries, or malnutrition. Moore et al90 conducted a comprehensive summation of their prolific research in this area, and concluded that MPS reached a plateau in younger men (early 20’s) with a protein dose of 0.24 g/kg BW, while in older men (early 70’s) MPS reached a - 62 - plateau after ingestion of 0.40 g/kg BW. Note that these are mean values (averages), so the upper 95% CIs (high-end limits) are worth pointing out, since individual response potentially can dictate requirements significantly above the mean. Quoting their discussion of this important nuance about protein doses that max-out MPS (my bolding for emphasis):90 “Additionally, it should be noted that the breakpoint observed in the present study would reflect the estimated average requirement to maximize MPS and, as such, the acute protein intake may be as high as ~0.60 g/kg for some older men (depending on the presence of potential contributing factors to the “anabolic resistance” of MPS) and ~0.40 g/kg for some younger men.” A notable example of a higher protein dosing ceiling for maximal MPS in older subjects than we previously thought, is a recent study by Park et al.91 In older untrained adults (mean age 69.3 years), 70 g protein from beef patties elicited greater MPS than 35 g in the non-trained state. Without an intermediate dose between 70 & 35 g, questions still remain about the precision of where beyond 30 g the anabolic ceiling was for this particular population. In contrast, Holwerda et al,92 examined the anabolic dose-response to 15, 30, and 45 g milk - 63 - protein concentrate taken post-resistance exercise by untrained older adults (mean age 66 years). Although wholebody protein synthesis was highest with the 45 g dose, MPS plateaued at 30 g. It’s tough to explain the discrepancy between these studies’ findings aside from the possibility that the mixed-macronutrient meal used in Park et al91 could have been more inherently anabolic than an isolated protein source. It’s also worth noting that anabolic resistance is not necessarily an inevitable fate. Breen and Phillips93 make a distinction between the “frail” elderly (who are afflicted by various illnesses) and the “well preserved” elderly (who are not significantly afflicted by age-related comorbidities, and are physically active). I prefer to call the latter “trained” elderly, since “well preserved” sounds too much like beef jerky. Along these lines, a systematic review by Shad et al94 included 24 studies containing 48 study arms (study groups). Overall, 18 study arms showed evidence of age-related anabolic resistance, 30 study arms did not. Importantly, when resistance exercise and protein/amino acid-based nutrition were combined, only 2 of the 10 study arms showed age-related anabolic resistance. It thus was concluded that the optimization of resistance exercise and protein or amino acid - 64 - consumption can produce comparable MPS responses between young (18-35 years) and older individuals (>55 years). Summing up • Age-related anabolic resistance is characterized by a diminished muscular sensitivity to protein feeding, requiring a higher protein dose to max-out MPS in older vs younger subjects. • Per the findings of Moore et al,90 older subjects max-out MPS at 0.4-0.6 g/kg per meal. Consuming this dose at least 3-4 times in the course of the day would ensure that the targeted total is hit. • Anabolic resistance is apparent in the general/untrained elderly population, and especially apparent in the frail elderly. However, this phenomenon does not necessarily apply to healthy older subjects, especially those who combine resistance training and proper protein feeding.93-95 Side-note: when Moore et al published their review,90 Macnaughton et al30 had not yet “shook up” the research world with their finding that 40 g elicited greater MPS than 20 g whey protein in young resistance-trained adults after high-volume (full-body) resistance exercise. It remains to be seen whether the same protocol in older subjects would nudge the protein dosing ceiling up even further. - 65 - Another side-note: Moore et al’s range for older subjects (0.40.6 g/kg)90 is nearly identical to the per-meal dosing range Brad Schoenfeld and I recommended in our paper on the topic of protein dosing for maximizing the per-meal anabolic effect (0.4-0.55 g/kg).32 Brad and I factored Macnaughton et al’s work30 (among several other lines of evidence) into our recommendations, hence our higher dosing suggestion than the lower doses traditionally thought to only apply to older subjects. - 66 - Chapter 17: How much protein do adolescents need? Adolescence has been loosely defined as the teenage years, but perhaps more accurately, it’s the timeframe spanning from the beginning of puberty, to the beginning of adulthood.96 This has been cited as the period between 10-19 years of age,96 but it’s also been referred to as 12-18 years of age.97 Adolescence is the second wave of rapid growth in the human life cycle, with the first wave occurring from infancy to early childhood.98 Daily energy requirements reflect the rapid growth that occurs during adolescence:99,100 DAILY ENERGY REQUIREMENTS OF ADOLESCENTS (PAL = physical activity level) 11-14 years Average PAL99 Heavy PAL100 15-18 years Average PAL99 Heavy PAL100 Males Females 2500 kcal 2475-3175 kcal 2200 kcal 2300-2725 kcal 3000 kcal 3450-3925 kcal 2200 kcal 2855-2875 kcal Current ‘authoritative’ recommendations of protein intakes for adolescents are lacking. They’re largely stuck in the RDA zone (9-13 years: 0.95 g/kg, 14-18 years: 0.85 g/kg).101 Refer to Chapter 3 for a discussion on the inadequacy of the RDA for most healthy, active populations. - 67 - Summing up: • The RDA for protein is inadequate for most healthy, active populations. Recent literature reviews100,102 have proposed that the protein needs of physically active adolescents should mirror the adult recommendations of the position stands of the authoritative organizations. • Recently, Berg102 proposed that adolescent athletes should consume 1.2-2.0 g/kg, which reflects the most recent joint position stand of the Academy of Nutrition and Dietetics, Dietitians of Canada, and the American College of Sports Medicine.18 • It should be noted that since the adolescent population is undergoing particularly rapid growth, a lower-end intake of 1.2 g/kg might not be adequate, as seen in IAAOdetermined protein requirements of children, which ranged 1.3-1.55 g/kg.103 • For the general adolescent population, I would default to the ISSN’s recommended intake for physically active adults (1.4-2.0 g/kg).17 • However, needs could be higher in adolescent athletes who go through periods of caloric restriction (i.e., for weight class-based sports, or to meet the body composition needs of particular competitive seasons or phases). In this case, Hector and Phillips’ recommendation of 1.6-2.4 g/kg for athletes in hypocaloric conditions seems warranted.36 - 68 - Chapter 18: How do sex differences influence protein requirements? In the history of “official” nutrition recommendations, none of the major/authoritative scientific organizations have specified different protein requirements for men and women. Based on the evidence as a whole, this lack of delineation is warranted. In the literature, there are flickers of difference, which I’ll touch upon. The main sex-based difference relevant to estimating protein needs is the higher proportion of body fat in women, which defaults to a lower proportion of lean mass. Total bodyweight-based protein estimation can therefore be skewed toward overestimation of women’s needs (I’ll get to practical solutions to this as we sum things up). Sex-based differences in muscle protein synthesis (MPS) across various populations could lead us to hypothesize that men and women need different amounts of total daily protein to maximize muscle growth or retention. However, this has not consistently been the case. A classic review by Tipton104 relayed several studies in the late1990’s and early 2000’s showing that between the sexes, there are no differences in whole-body protein synthesis or - 69 - breakdown, and no difference in basal level of net muscle protein balance. Furthermore, there’s no human (or animal) evidence that ovarian hormones inhibit muscle protein synthesis. Subsequent research by Smith et al105 found no meaningful differences in MPS during fasting conditions and during hyperinsulinemia-hyperaminoacidema between young and middle-aged adult men and women. Continuing the same theme, Dryer et al106 found that post-exercise increases in MPS and mTOR signaling in leg muscle is not different between men and women. A recent (2018), thoroughly done review by Witard et al107 reported that sex-specific differences in MPS response to exercise and nutrition has not been consistently shown in young adults, but in older adults, greater basal MPS rates have been shown in older (>65 years). This can at least partially explain their slower age-related loss of muscle compared to older men.108 More recent data using the indicator amino acid oxidation (IAAO) technique found that resistance-trained men required 2.0-2.38 g/kg (2.2-2.6 g/kg FFM)26 and resistance-trained women required 1.49-1.93 g/kg (1.8-2.2 g/kg FFM),109 to maximize the acute anabolic response (these ranges represent - 70 - estimated average requirements and upper confidence limits). However, the main limitation of the IAAO method is that it assesses whole-body protein synthesis, not muscle-specific protein synthesis. Summing up • Overall, there’s insufficient evidence to program different protein targets either in total or per meal, based on sex. This lack of meaningful difference in anabolic response to protein feeding has been seen in the majority of studies examining this question. • While younger adults consistently fail to exhibit sex-based differences in MPS response to nutrition and training, it’s possible that the greater basal/resting MPS rates in older women (>65 years) can explain their slower rate of muscle loss. However, this is largely speculative, since a multitude of genetic and lifestyle factors are involved, and can vary widely across individuals. • Keeping in mind that with an optimal total daily protein intake of 1.6-2.2 g/kg, both sexes can use 1.6 g/kg as a baseline from which to adjust upward. It’s reasonable to assume that women require less protein per unit of total body mass, since they tend to carry a higher proportion of - 71 - body fat than men. Therefore, women’s needs along the 1.62.2 g/kg range will likely hover around the lower rather than the higher end of the range. • Fat-free mass (FFM)-based protein targets are more complicated and not necessarily more accurate, but they can satisfy some people’s OCD. 2.0 g/kg FFM (0.9 g/lb FFM) is a reasonable baseline target for general purposes. Individuals specifically aiming for muscle gain (as well as dieters running a caloric deficit) can start at 2.2 g/kg FFM (1.0 g/lb FFM), and adjust according to individual response. - 72 - Chapter 19: How do high-protein diets impact bone and kidney health? Bone health Bone health and kidney function are perhaps the most common concerns surrounding high protein intakes. We’ll start with bone. Who hasn’t heard the claim that protein, which is made of amino acids, leeches calcium from bone in order to buffer the acidity of a high-protein intake, which eventually leads to bone loss. Well, that sounds logical on paper, but it doesn’t stand the test of science. The following excerpt from a classic review by Layman5 is worth quoting because it eloquently nails the important points: “The efficacy of calcium and protein are interrelated. Calcium supplements are largely ineffective for remodeling of bone matrix if protein is limiting. Positive effects of calcium appear to require intakes of protein >1.2 g/kg to have beneficial effects. The long-held belief that increased dietary protein could cause bone loss as reflected in increased urinary calcium is incorrect, and protein is now recognized to increase intestinal calcium absorption in addition to enhancing bone matrix turnover.” - 73 - More recently, Antonio et al110 found that a high protein intake (averaging 2.8 g/kg) over a 6-month period in trained women had no harmful effects on bone mineral content or density. Subsequently, a meta-analysis by Groenendijk et al111 showed that high (above the RDA of 0.8 g/kg) vs low protein intake resulted in a significant decrease in hip fractures. Kidney function Antonio et al conducted a series of investigations on resistancetrained subjects refuting the claim that high protein intake is an inherent threat to kidney function.112-114 An 8-week trial involving a protein consumption at 3.4 g/kg showed no adverse effects on renal function (or in general).104 Subsequently, a 4-month trial ranging 2.6-3.3 g/kg showed no adverse clinical effects, including renal function.113 Finally, a 1year crossover trial involving 2.5 & 3.3 g/kg (6 months at each protein level) showed no harmful effects on any biomarkers of health, including liver and kidney function.114 A recent systematic review and meta-analysis by Devries et al115 involving 28 studies and 1358 subjects compared highprotein intakes (≥1.5 g/kg body weight or ≥20% energy intake or ≥100 g protein/day) with normal/lower-protein intakes - 74 - (≥5% less energy intake from protein/day compared with high-protein group) on kidney function. It was concluded that higher protein intakes have a trivial to non-existent effect on kidney function. Summing up: • Current evidence does not support concerns about high protein intakes being inherent threats to kidney and bone of healthy individuals. • Protein intakes above the RDA (0.8 g/kg) are associated with a decrease in hip fractures, and intakes below 1.2 g/kg might not optimize calcium metabolism as it relates to bone health. • Protein intakes as high as 2.8 g/kg for 6 months have not shown adverse effects on bone. • Protein intakes as high as 3.4 g/kg have not shown any adverse effects on kidney function in the intermediate-term, and intakes at 2.5 & 3.3 g/kg have not shown any adverse effects on kidney function in the long-term. • If you have (or suspect you might have) kidney disease, the research discussed in this chapter might not apply to you. Consult with your doctor (and if possible, your renal dietitian) if you have any doubts or concerns. - 75 - Chapter 20: Is there an inherent advantage to pre-bed casein? Cow’s milk protein consists of 80% casein and 20% whey. Whey is the fast-digesting protein. Casein is a slow-digesting protein, which gives it the potential for special application at the pre-bed time point, for the purpose of a timed-release of amino acid availability. Pre-bed protein increases amino acid availability through the night, thus resulting in net gains in muscle protein balance compared to…well, no protein feeding at all.28 In light of the past several years of pre-bed protein research, a recent review by Kim116 concluded that 40-48 g casein ingested roughly 30 minutes prior to sleep enhances the anabolic response in skeletal muscle – especially when this is done in the post-exercise state. While the above conclusion is reasonable based on the existing findings, a potentially more practical question is whether the timing of casein intake (pre-bed versus some other point in the day) makes a difference on muscle growth or muscle retention in the context of a resistance training program and optimized total daily protein intake. The answer to this would require longitudinal research (lasting several weeks or months. To - 76 - date, there are only two studies that directly examine this question, which I’ll discuss next. Antonio et al117 found no significant effect as a result of 54 grams of casein either in the morning (before 12:00 noon) or evening (90 minutes prior to going to bed) added to the habitual routine of resistance-trained subjects in an 8-week period. No between-group or within-group differences were seen, as if the 54 g casein disappeared into thin air. The morning & pre-bed casein groups had a baseline/habitual protein intake of 1.7 and 1.9 g/kg respectively. It’s possible that the additional protein was superfluous in the face of protein intakes that were already optimized. Casein supplementation brought protein intakes up to a total of 2.4 g/kg in both groups. A subsequent study by Joy et al118 on recreationally trained subjects compared the a 10-week effects of 35 g casein either in the morning (with 35 g maltodextrin taken pre-bed) or prebed (with 35 g maltodextrin taken earlier in the day). Baseline protein intakes of the morning and pre-bed casein groups were not reported, but intake during the trial (including casein supplementation) was 1.99 g/kg and 2.14 g/kg, respectively. Both groups had significant increases in muscle strength and mass, with no significant differences between groups. - 77 - Summing up The scant evidence to date does not show that specifically timing casein before bed imparts special effects on body composition compared to casein ingestion elsewhere in the day – particularly in resistance trainees consuming optimal total daily protein amounts (approximately 1.6-2.2 g/kg24). If there is an advantage, which in theory would be greater net gains in muscle protein balance leading to greater muscle gain or retention, it’s been too small to detect within the methodological parameters of the studies thus far. It would be interesting to see future studies compare the body composition effects of casein with different proteins taken prebed, to see if casein is actually something special at that time period. Again, this would ideally be done via parallel arm study involving resistance training, drawn out for several weeks. This is not likely to happen any time soon, but if/when it does, we’ll be ready. :) - 78 - Chapter 21: How does postworkout whey compare to chicken, beef, or casein for improving body composition & strength? The whey, chicken, & beef study This question is relevant in light of the enduring interest in post-exercise tactics that might enhance muscular adaptations to resistance training. Whey has been studied extensively for this purpose. Direct comparisons with other (non-milk-based) animal proteins have been elusive – until 2017. An 8-week study by Sharp et al119 compared the effects 46 g whey, chicken, or beef protein post-exercise in resistance-trained, collegeaged men and women. A fourth group served as a control, taking a maltodextrin-based placebo. All three protein groups significantly increased lean body mass, and significantly decreased fat mass, with no significant differences between groups. The control group also showed recomposition improvements, but not to a statistically significant degree. The greater LBM gains in the protein groups compared to the control group are interesting, since their dietary protein intakes were reported to range 2.1-2.2 g/kg while the placebo group’s protein intake was 2.0 g/kg. All of these intakes fall well - 79 - into the realm of optimal protein dosing. In the strength department, 1-rep maximum for deadlift and bench press significantly increased compared to baseline for all groups, including the control group. No significant differences between groups were detected. Keep in mind, the chicken and beef protein hydrolysate powders were used instead of these proteins in their native/flesh form. Such a comparison would be messier and more logistically challenging, but more relevant to dietary programming in the real-world. What about casein vs whey? Whey’s stimulation of muscle protein synthesis (MPS) is a steeper/sharper spike, while casein’s MPS curve is flatter and longer. However, acute (short-term response) studies comparing these proteins have consistently shown no difference in MPS120,121 or net protein balance122 when ingested post-exercise. Longitudinal studies measuring changes in body composition and strength performance are mixed, with one study showing the superiority of whey123 and two showing no significant - 80 - difference between whey and casein ingested pre- and/or postexercise.124,125 The one study showing whey as the winner123 had a dosing protocol that makes me do a double-take every time (1.5 g/kg; 90 g per day of the protein supplements, in addition to their habitual diets). This study also has a higher potential for bias since the lead investigator was also the research director of the company that provided the whey protein product used in the study. This obviously doesn’t guarantee bias, but it still raises my skeptical senses. Don’t get me wrong – I love how it validates my own high intake of whey, but hard not to see this study as an oddity in the literature. Summing up The collective evidence thus far does not indicate that the specific postworkout protein source makes a significant difference in the enhancement of resistance training adaptations – especially in the context of a diet containing a mix of high-quality proteins. - 81 - Chapter 22: How do whole eggs compare with egg whites for muscle growth? Acute anabolic response van Vliet et al126 found that in healthy, resistance-trained men, the post-exercise consumption of 3 whole eggs resulted in a 29.4% greater stimulation of muscle protein synthesis (MPS) than 6 egg whites (a protein-equated dose). Short-term MPS response is useful for investigating mechanism, but it’s hypothesis-generating until things are put to the test in longitudinal research capable of measuring changes in body composition. Fortunately, this has been done. Let’s have a look. Adaptations to training Bagheri et al127 compared the 12-week effects of 3 whole eggs vs 6 egg whites ingested post-exercise in resistance-trained men undergoing a nonlinear, periodized, progressive resistance training program. Total bodyweight gain in the whole egg & egg white group was 1.7 & 1.8 kg, respectively, indicating that a similar net caloric surplus was sustained in both groups. Total daily protein intake in the whole egg & egg white groups was 1.42 & 1.47 g/kg, respectively. This leaves - 82 - open questions about whether significant differences would be seen with optimized protein intakes of 1.6-2.2 g/kg.24 Lean mass gain was greater in the whole egg group (3.7 kg vs 2.9 kg), but not to a degree of statistical significance. Fat mass decreased in both groups (2.0 & 1.1 kg in the whole egg & egg white group, respectively), but this difference did not reach statistical significance. However, body fat percentage decrease was significantly greater in the whole egg group. Total protein intakes in the whole egg & egg white group were respectively. In the strength department, whole eggs resulted in significantly greater gains in knee extension & handgrip strength. Another factor working in the favor of whole eggs’ anabolic potential is their significantly greater impact on serum testosterone levels, which increased by 240 & 70 ng/dL in the whole egg & egg white group, respectively. The clinical significance of this testosterone hike is up for debate, but could plausibly function as part of an anabolic mechanism beyond the extra calories from the fat content of the yolks. Summing up: The scant body of research in this area shows that whole eggs have greater acute and chronic anabolic effects than a protein- 83 - equated dose of egg whites consumed post-exercise. An additional effect of whole egg consumption is an increase in testosterone. I wouldn’t say that egg yolks can make or break body composition goals. It’s a relatively minor variable within the multitude of programming elements that facilitate muscle growth. However, based on the evidence thus far, including egg yolks in the diet might provide a slight edge compared to eating just the whites. The data on eggs and health outcomes are a mixed, equivocal mess that’s difficult to distill into definitive recommendations.128 However, considerable evidence has mounted in defense of eggs, challenging traditional concerns and caveats about whole egg consumption..129 If you’re worried enough, keep regular track of your bloodwork to note any changes (good or bad) that might correlate with your egg intake, and heed the advice of your doctor. - 84 - Chapter 23: If collagen is considered a lowquality protein, does that make collagen supplements useless? Collagen is the most abundant protein in the entire animal kingdom.130 It’s also the most abundant protein in humans (and most vertebrates), comprising up to a third of total bodily protein mass.131 There are 16 types of collagen in the body, 8090% of which are types 1-3. The different collagen types and their characteristics are tabulated here.130 However, anyone who’s taken a basic nutrition course was taught that collagen is the one animal protein source that’s comparatively inferior. Collagen has a low indispensable-todispensable amino acid ratio, and is devoid of the essential amino acid tryptophan, which has led to the classification of collagen as an "incomplete" protein.132 However, the high glycine, proline, and hydroxyproline content of collagen makes it functionally unique among other proteins. And as the saying goes, that’s where the magic happens. Summing up: • The current evidence supports the potential of supplemental collagen for strengthening ligaments & tendons,133,134 mitigating osteoarthritis & osteoporosis,135,136 reduced- 85 - • • • • activity joint pain,137,138 and dermatological applications.139,140 Although collagen supplementation has been shown to increase lean body mass and strength in older141,142 and younger adults,143 these studies did not directly compare collagen with another protein source. In direct comparisons, whey has outperformed collagen for decreasing android fat144 and increasing acute & longer-term muscle protein synthesis.145 The effective dosing ranges 8-15 g.135,141-143 Increased plasma glycine and proline levels have been reached with 8 g/day. 12g/day improved symptoms of osteoarthritis & osteoporosis. 15 g/day resulted in lean mass & strength gains (note that these results were not from direct comparisons to other proteins). - 86 - Chapter 24: How do plant proteins compare to animal proteins for muscle growth? The popularity of plant-based diets has been on the rise as of roughly the past decade.146 Plant proteins have long been considered inferior to animal proteins due to their lower digestibility & lesser proportion of essential amino acids (EAA), including key anabolic drivers, the branched-chain amino acids; leucine in particular.147 Nevertheless, claims of plantbased proteins being on a level playing field with animal-based proteins for supporting muscle growth have been gaining momentum in recent years, so let’s have a look at the evidence. Short-term anabolic response Whey has outperformed soy in acute muscle protein synthesis (MPS) studies in both young148 and old subjects.149 And frankly, these findings were not surprising, given the higher EAA content of whey. The acute anabolic response superiority of whey has also been seen in direct comparisons to wheat protein,150 as well as pea-based plant protein blends.151 Monteyne et al152 recently reported that a fungus-derived protein product (mycoprotein) outperformed milk protein for raising MPS.23 However, the greater amount of protein (31.5 vs 26.2 g) and kcal (238 vs 108) in the mycoprotein treatment - 87 - confounds the comparison. More recently, Pinckaers et al,153 similar elevations in MPS were seen in a comparison of corn protein, milk protein, and a corn-milk protein blend. Chronic effect studies In 2013, Joy et al154 compared post-exercise whey and rice protein isolate supplementation in resistance-trained subjects. Both treatments similarly improved body composition and exercise performance. Babault et al155 shook up the protein world in 2015 when they published findings showing the superiority of pea protein over whey for increasing muscle thickness in untrained subjects. Of course these results are intriguing, but I’d view this finding with caution until this study is replicated. Nieman et al156 recently reported that whey outperformed pea protein at suppressing blood biomarkers of muscle damage from 5 days of eccentric-based exercise. A 2018 meta-analysis by Messina et al157 examined the effect of animal protein versus soy protein supplementation on muscle size and strength gains in subjects undergoing resistance training. Of the 9 studies, 3 favored dairy protein, and 6 studies showed no significant advantage of either protein type. None of the studies actually found soy to be the superior performer. - 88 - A recent 12-week study by Hevia-Larraín et al158 is the first to compare the effects of a completely plant-based (vegan) diet with an omnivorous diet on the resistance training adaptations, under conditions of optimal protein intake (at least 1.6 g/kg). Both groups showed increased lean mass, cross-sectional area of type 1 & 2 fibers, and leg press strength, with no differences between groups. Body fat was unchanged compared to baseline in both groups. Protein supplementation in the omnivores and vegans were in the form of whey (41 g/day) and soy protein (58 g/day), respectively. While the lack of differences is interesting, the use of untrained subjects is the Achilles’ heel that leaves open questions. In newbies, potential advantages in one in treatment versus the other are often masked by an indiscriminately robust response to whatever novel protocol is undertaken. Replication of this study with trained subjects would provide useful data. Summing up • In the majority of acute (short-term) anabolic response studies to date, commonly used animal proteins (dairy-based proteins in particular) have outperformed commonly used plant-based proteins. • Longitudinal studies capable of measuring effects on body composition and exercise performance have been mixed, - 89 - with no clear advantage of animal- versus plant-based protein supplementation. • Only one study to date158 has compared plant versus animalbased protein supplementation in the context of a completely plant-based (vegan) diet versus omnivorous diet, and no between-group differences were seen in lean mass & strength gains. A caveat is the use of untrained individuals, who are subject to the newbie gains, which can mask otherwise detectable differences in the treatments compared. • In Hevia-Larraín et al’s study,158 protein supplementation was used to facilitate an optimal daily intake. The vegan group supplemented with soy protein isolate (58 g/day). • On a related note: concerns about the isoflavone content of soy foods having feminizing effects in men are not substantiated by the literature as a whole.159 However, this concern is not completely unfounded, since testosterone decreases have been reported with soy protein isolate supplementation dosed at 20 g/day160 and 56 g/day.161 Limitations of this research aside, sole reliance on large amounts of soy protein supplementation to hit daily protein targets is probably a not good idea for those concerned with hormonal effects. - 90 - • Animal proteins possess an anabolic advantage (and in some cases an ergogenic advantage) due to higher EAA content – particularly leucine, and also due to constituents such as taurine, carnosine, creatine,162,163 collagen,132-143 and even cholesterol,127,164,165 none of which are present in plant foods. • While it’s possible for exclusively plant-based protein intake to perform similarly to omnivorous protein intake when enough total protein is consumed (≥1.6 g/kg), animal-based protein is generally more anabolic on a gram-for-gram basis. In support of this point, I’ll end off with a salient excerpt from a recent review by Berrazaga et al:147 “Despite some contradictions, taken together, most of these studies suggest that the difference between the anabolic effects of plant- and animal-based proteins could be reduced with an adequate (i.e., increased) protein intake. […] Nevertheless, at similar protein intakes, most studies have reported a lower ability of plant-based protein sources to stimulate protein synthesis at the skeletal muscle level and induce muscle mass gain compared to animal-based protein sources, especially in older people. The lower anabolic effect of plant-based protein sources is partly due to their lower digestibility and their lower essential amino acid content, especially leucine, compared to animal proteins.” - 91 - Chapter 25: How satiating is protein, really? Protein is considered to be the most satiating macronutrient.166,167 This is based on an abundance of evidence from short-term satiety response data, as well as longitudinal studies comparing higher versus lower protein intake on bodyweight and body composition. Before getting into that, let’s pay a quick tribute to accuracy by getting the definitions straight. As Bellisle et al168 eloquently put it: “Satiation occurs during an eating episode and brings it to an end. Satiety starts after the end of eating and prevents further eating before the return of hunger.” Most acute (short-term) fixed-meal studies have shown significant reductions in hunger and increased fullness after consuming higher versus standard protein meals.169 The most consistent hormonal responses associated with higher-protein meals are increases in the ‘satiety hormones’ PYY and GLP-1. These acute findings have been reflected in longer-term studies consistently showing the superiority of higher protein intakes (1.2-1.6 g/kg or ≥25% of total kcal) compared to lower intakes on weight/fat loss, weight loss maintenance, and lean mass preservation.169-171 - 92 - A possible limit to protein dosing for satiety? A recent 7-day crossover trial by Roberts et al172 compared the satiating effect of a moderate-protein diet (1.8 g/kg) with a high-protein diet (2.9 g/kg) in resistance-trained subjects in hypocaloric conditions (20% below maintenance needs). This 7-day hypocaloric period was followed by a 3-day ad libitum (unrestricted) period. Habitual training routines were maintained throughout the study. There was a lack of difference in satiety between the two groups in most of the parameters tested. However, it’s notable that during the hypocaloric conditions, 2 of the 9 questionnaire items favored the high-protein diet. Specifically, subjects reported increased cravings on the moderate-protein diet, and greater satisfaction on the highprotein diet. The authors concluded that in sum, there was insufficient justification for recommending the high-protein (2.9 g/kg) over the moderate-protein intake (1.8 g/kg) for the pursuit of maximizing satiety in hypocaloric conditions, with the exception of implementing high-protein meals to mitigate cravings if needed. I think this study is interesting, and provides some good food for thought. However, the 7-day period leaves a lot of unanswered questions, especially due to - 93 - the appearance of a trend favoring the satiating capacity of the higher-protein intake. A series of original investigations by Antonio et al112-114,173 spanning 8 weeks to 6 months on resistance-trained subjects involved baseline protein intakes of ~2.0-2.2 g/kg that were increased to 3.3-4.4 g/kg with no significant changes in body composition overall (except one study saw greater recomposition in the high-protein condition112). It’s remarkable that in free-living conditions, increasing baseline protein intakes by approximately 50-100% for months did not result in significant weight or fat gain. It’s likely that heightened satiety drive down the intake of the other macronutrients, since the ‘disappearance’ of the surplus protein calories cannot all be explained by increased thermogenesis (energy expenditure). Summing up Protein is the king of the macronutrients for satiety. In hypocaloric conditions, it’s debatable whether 1.8 g/kg (vs 2.9 g/kg) is sufficient to max-out satiety, although there seems to be an advantage for suppressing cravings and increasing satisfaction with 2.9 g/kg.172 In the case of protein overfeeding in resistance trainees, there’s potentially some satiety magic with intakes at or beyond approximately 3 g/kg.112-114,173 - 94 - Chapter 26: What about non-linear protein intake through the week (training days vs nontraining days, protein hyperfeeds)? Carb-cycling, or non-linear carbohydrate intake through the course of the week has been a perennial topic at the forefront of physique and athletic performance pursuits. Less on-theradar is non-linear protein intake through the course of the week, and how it might benefit body composition goals. Training days vs. non-training days A question I’ve been frequently asked is whether protein needs differ on training versus non-training days. I answer most questions with a couple of questions of my own: What’s the goal, and what are the stakes? That’s my way of getting the people to think about what they want to accomplish, what their current training &/or body comp status is, and how much is riding on the attainment of this goal. There’s a wide range of possible scenarios to consider. To simplify things for this discussion, we can focus on two general targets: the goal of maximizing muscle gain, and the goal of muscle retention while dieting. Maximally facilitating muscle gain involves sustaining a net caloric surplus.22 Given this, it’s - 95 - not typical for people seeking mainly to gain muscle to search for ways to economize their caloric intake. There is a segment of the protein research audience concerned with the potential for high protein intakes to negatively impact longevity. However, as discussed in Chapter 15, this is a far-fetched idea. Nevertheless, I’m still regularly asked if it’s necessary or beneficial to lower protein intake on non-training days. This would potentially be warranted if the training-induced elevation in muscle protein turnover (synthesis & breakdown) was a very short-lived phenomenon. This is not the case. Protein turnover can stay elevated for 24-48 hours after a bout of resistance exercise.174-176 Burd et al177 found that myofibrillar MPS and anabolic signaling were enhanced by protein feedings for at least 24 hours after a bout of resistance exercise. Damas et al178 examined muscle protein changes in the 24-48 post-exercise period, and found that muscle hypertrophy occurs via gains in myofibrillar MPS mainly after the attenuation (lowering) of muscle damage. This reinforces the importance of optimizing dietary protein intake to maintain positive net balance of muscle protein turnover. The main message here is that enhanced muscle sensitivity and receptivity to protein/amino acids persists far beyond the immediate post-exercise period, and well into the following - 96 - day (and possibly beyond). Thus, for the goal of maximal utilization of protein during the 24-48-hour recovery period where pushing for positive net protein balance is crucial for muscle growth, low-balling protein intake during non-training days is a bad idea. The what, why, & how of protein hyperfeeds “Protein hyperfeed” is a cool-sounding term I came up with. It’s the protein equivalent of a carb-up without the low intake part of the cycle. While carb refeeds are large bouts of carb intake alternated with carb restriction of varying severity, protein hyperfeeds are high-protein days (2-2.5 times the normal intake) placed strategically through the week. “Normal” protein intake would actually be defined as what’s optimal in normal circumstances (≥1.6 g/kg per day). Hyperfeeds would put protein at approximately 3.2-4.5 g/kg, or 1.45-2.0 g/lb for the day. For example, a 75 kg person might consume 135 g protein on a normal day. A hyperfeed day would provide 2-2.5 times this amount, which is 270-337 g. There are three main uses/objectives for protein hyperfeeds that can be pursued separately or in combination, depending on the individual’s goals and current training status: 1) to maximize satiety and minimize cravings in general, and - 97 - especially on carb-restricted days, 2) to allow bouts of ad libitum/unrestricted eating for the purpose of alleviating psychological “diet fatigue” with minimal risk for undue fat gain, and 3) to push toward recomposition in higherintermediates and advanced trainees. This third point is more of a dice-roll. Nevertheless, in my observations, even when protein hyperfeeds are calculated as net surplus calories, there is still a “disappearance” of the extra protein, as reported repeatedly by Antonio et al,112-114,173 whose subjects saw either no significant change, or favorable change in body composition (fat loss and/or muscle gain) as a result of no other program change aside from increasing protein intakes to ≥3 g/kg. Potential explanations for Antonio et al’s “disappearing” protein surpluses include increased thermogenesis (both dietary, exercise-based, and non-exercise-based), increased satiety driving down the intake of the other macronutrients, misreporting of intake, and increased excretory energy losses.33 I witnessed this type of protein-induced recomp in clients several years before Antonio et al’s protein overfeeding studies, so while some viewed the findings with skepticism, they actually reflected my notes from the trenches. In eucaloric (maintenance) conditions, and even in purposefully targeted - 98 - net weekly surplus conditions, I have observed recomp in clients on regular protein hyperfeeds. There are a multitude of variables that make recomp possible,44 but in my observations, experimenting with protein hyperfeeds through the week is a tool worth trying out for this purpose. In my experience and observations in clients, 2-3 protein hyperfeeds per week is what most people can tolerate (and enjoy). Protein hyperfeeds should ideally be positioned on, or immediately before days where cravings are historically highest for the individual. Or, choose any day(s) where carbohydrate intake is not elevated for refeeding or carbing-up. An interesting phenomenon I’ve observed repeatedly is that the satiating effects of the protein hyperfeed can last through the entire following day. Individual responses vary here. Summing up • Rather than low-balling protein on days off, it’s best to keep protein at levels known to optimize adaptations to resistance training. Protein turnover and heightened muscular sensitivity to protein feeding can last 24-48 hours after a training bout. • Protein hyperfeeds are a tool I’ve used with dieters seeking novel tactics to increase satiety and minimize the - 99 - psychological fatigue of dieting. It has also worked for intermediate and advanced trainees seeking recomposition. • My field experience with protein hyperfeeds reflects the intriguing results of Antonio et al’s protein overfeeding studies,112-114,173 as well as Bararat et al’s recent review reporting that a protein intake of 2.6-3.5 g/kg of FFM is associated with recomposition.44 The protein hyperfeed protocol I’ve found success with involves 2-2.5 times normal protein intake, 2-3 days per week. Protein sources should be kept lean, for the most part. • Protein hyperfeeds also make protein lovers extremely excited about their diets - especially those who prefer more savory foods than sweet foods. Those with more of a sweet tooth can still engage in protein hyperfeeds, since endless dessert variants can be derived from protein powder. • Who knows, maybe there’s some metabolic magic to be had from protein hyperfeeds. We’ve got the insulin fairy, might as well add the protein fairy to the mix (to my newer readers who might not get that, it’s a corny inside-joke for those who have been following my work since the olden days). - 100 - Chapter 27: Protein servings & sources Categorizing foods is a messy endeavor since there’s inevitable overlap in macronutrition and micronutrition. Nevertheless, a general awareness of the fat levels of your protein sources can help you make the right choices based on the nature of the overall macronutrient profile you’re aiming for. A note about the listed macronutrient values of each food: they almost never add up exactly to the total calories per serving. This is an unavoidable shortcoming. I used nutritiondata.com, for the most part. It relies on the USDA's National Nutrient Database. Even the most sophisticated and reliable databases are unable to list nutrient stats with complete accuracy. This is partially due to the necessity of rounding off values. However, the important thing is that the numbers are close enough. Thinking you’ll hit everything perfect down to the gram is not just unrealistic, it’s also an unnecessary level of micromanagement that will not make or break anyone’s results. The following food lists are separated into animal and plantbased sources. Leucine content is included. With few exceptions such as protein powder, plant-based protein sources have a lower proportion of protein and higher proportion of carbohydrate and/or fat, so they pack less protein per calorie. - 101 - Remember that serving sizes are arbitrary (& adjustable) I listed serving sizes that (for the most part) hit 20 g protein or more. You’ll notice that many of the plant-based servings come up shy of 20 g protein. However, you can adjust all of the serving sizes (or number of servings) up or down to hit a particular protein dose. Just be aware of how serving adjustments impact total calories. On a final note, keep in mind that this is not a comprehensive list of foods. The aim is to give you a quick reference, and help you get some ideas. VERY LEAN PROTEIN SOURCES: ANIMAL-BASED Food Serving size Beef, ground, 95% lean 3.5 oz meat (100 g) Beef, top sirloin, separable 3.5 oz lean only, trimmed to 0" (100 g) fat, select, cooked, broiled Casein protein powder 1 scoop (combination of micellar (32 g) casein and milk protein concentrate), Catfish, cooked, dry heat 3.5 oz (100 g) Cheese, cottage, 2% 1 cup milkfat (238 g) Cheese, low-fat, cheddar 2/3 cup, or colby shredded (75 g) Chicken breast, meat only, 3.5 oz roasted (100 g) kcal 171 Prot (g) 26.3 Carb (g) 0 Fat (g) 6.5 Leu (g) 2.0 177 30.8 0 5.0 2.5 120 24.0 5.0 1.0 2.0 105 18.5 0 2.8 1.5 180 22.0 6.0 5.0 2.2 129 18.1 1.5 5.2 1.8 165 31.0 0 3.6 2.3 - 102 - VERY LEAN PROTEIN SOURCES: ANIMAL-BASED Food Clam, mixed species, cooked, moist heat Cod, Atlantic, cooked, dry heat Egg whites, raw, fresh Orange roughy, cooked, dry heat Pork, fresh, loin, tenderloin, fat-trimmed, roasted Salmon, canned, solids with bone and liquid Sea bass, mixed species, cooked, dry heat Shrimp, mixed species, cooked, moist heat Snapper, mixed species, cooked, dry heat Tuna, light, canned in water, drained solids Turkey breast, skinless, cooked, roasted Trout, rainbow, cooked, dry heat Whey protein powder (mix of isolate and concentrate) Whitefish, mixed species, cooked, dry heat Yogurt, Greek style, nonfat Serving size 3.5 oz (100 g) 3.5 oz (100 g) 1 cup or 8-12 whites (243 g) 3.5 oz (100 g) 3.5 oz (100 g) kcal 148 Prot (g) 25.5 Carb (g) 5.1 Fat (g) 1.9 Leu (g) 1.8 105 22.8 0 0.9 1.8 117 26.5 1.8 0.4 2.5 105 22.6 0 0.9 1.8 143 26.2 0 3.5 2.2 3.5 oz (100 g) 3.5 oz (100 g) 3.5 oz (100 g) 3.5 oz (100 g) 3.5 oz (100 g) 3.5 oz (100 g) 3.5 oz (100 g) 1 scoop (32 g) 3.5 oz (100 g) 1 cup (224 g) 139 19.8 0 6.1 1.6 124 23.6 0 2.6 1.9 99 20.9 0 1.1 1.7 128 26.3 0 1.7 2.1 116 25.5 0 0.8 2.1 135 30.1 0 0.7 2.4 169 24.3 0 7.2 1.9 120 24.0 3.0 1.5 2.5 172 24.5 0 7.5 2.0 120 20.0 9.0 0 1.6 - 103 - LEAN TO MODERATE-FAT PROTEIN SOURCES: ANIMAL-BASED Food Serving size 3.5 oz (100 g) 3.5 oz (100 g) kcal 2/3 cup (150 g) Cheese, mozzarella, part 3.5 oz skim milk (100 g) Cheese, ricotta, part skim 2/3 cup milk (162 g) Chicken, leg, meat and 3.5 oz skin, roasted (100 g) Duck, domesticated, meat 3.5 oz only, cooked, roasted (100 g) Egg, whole, hard-boiled or 3 large poached (150 g) Lamb, foreshank, trimmed 3.5 oz to 1/8" fat, braised (100 g) Pork, fresh, loin, center rib 3.5 oz (chops), bone-in, separable (100 g) lean and fat, braised Salmon, Atlantic, cooked, 3.5 oz dry heat (100 g) Sardines, canned in tomato 3.5 oz sauce, drained solids with (100 g) bone Turkey sausage, fresh, 3.5 oz cooked (100 g) Beef, ground sirloin, 90% lean meat Beef, ground, 85% lean meat/15% fat, patty, broiled Cheese, feta, reduced fat 214 Prot (g) 26.6 Carb (g) 0 Fat (g) 11.1 Leu (g) 2.0 250 25.9 0 15.5 2.0 206 21.0 0 13.0 1.7 254 24.3 2.8 15.9 2.4 224 18.5 8.3 12.9 2.0 232 26.0 0 13.5 1.9 201 23.5 0 11.2 2.0 232 18.9 1.8 15.9 1.6 243 28.4 0 13.5 2.2 250 26.7 0 15.1 2.1 206 22.1 0 12.3 1.8 186 20.9 0.7 10.5 1.4 196 23.9 0 10.4 1.7 - 104 - HIGH-FAT PROTEIN SOURCES: ANIMAL-BASED Food Serving size Bacon 2 oz (56 g) Beef, corned beef, brisket, 3.5 oz cooked (100 g) Beef sausage, fresh, cooked 3.5 oz (100 g) Beef tongue, cooked 3.5 oz (100 g) Cheese, American 3 oz (85 g) Cheese, blue 3 oz (85 g) Cheese, cheddar 2/3 cup, (75 g) Cheese, feta 1 cup (150 g) Cheese, goat 3 oz (85 g) Cheese, Monterey jack 3 oz (85 g) Cheese, Swiss 3 oz (85 g) Sausage, beef, fresh, cooked 3.5 oz (100 g) Sausage. pork, fresh, cooked 3.5 oz (100 g) kcal 298 Prot (g) 21.4 Carb (g) 0.8 Fat (g) 22.6 Leu (g) 1.7 251 18.2 0 19.0 1.4 332 18.2 0 28.0 1.5 284 19.3 0 22.3 2.0 315 18.6 1.2 26.4 1.6 297 18.0 2.1 24.0 1.6 300 18.5 0.9 24.8 1.6 396 21.3 0 31.9 1.7 381 25.5 1.8 30.0 2.1 300 21.0 1.0 27.0 1.6 318 22.5 4.5 23.4 1.8 332 18.2 0.4 28.0 1.5 339 19.4 0 28.4 1.3 - 105 - VERY LEAN PROTEIN SOURCES: PLANT-BASED Food Beans, black, cooked Beans, kidney, all types, cooked Beans, lima, large, cooked Beans, navy, cooked Beans, pinto, cooked Beans, white, cooked Beans, refried, canned Serving size 1 cup (172 g) kcal 1 cup (177 g) 1 cup (188 g) 1 cup (182 g) 1 cup (171 g) 1 cup (179 g) 1 cup (238 g) 1 cup (164 g) 1 cup (172 g) Chickpeas (garbanzo beans, cooked Cowpeas, common (blackeyes, crowder, southern), cooked Edamame (soybean), frozen, 1 cup prepared (155 g) Lentils, cooked 1 cup (198 g) Peas, green, cooked, boiled, 1.5 cups drained (240 g) Pea protein isolate 1 scoop (28 g) Soy protein isolate 1 scoop (28 g) Veggie burgers or soy 2 patties burgers (140 g total) 227 Prot (g) 15.2 Carb (g) 40.8 Fat (g) 0.9 Leu (g) 1.2 225 15.3 40.4 0.9 1.3 216 14.7 39.3 0.7 1.3 255 15.0 47.8 1.1 1.3 245 15.4 44.8 1.1 1.1 249 17.4 44.9 0.6 1.4 217 12.9 36.3 2.8 1.0 269 14.5 45.0 4.2 1.0 200 13.3 35.7 0.9 1.0 189 16.9 15.8 8.1 1.2 230 17.9 39.9 0.8 1.3 187 12.3 34.2 0.6 0.7 110 23.1 1.7 1.7 1.9 94.6 22.6 2.1 0.9 1.9 248 22.0 20.0 8.8 1.9 - 106 - LEAN TO MODERATE-FAT PROTEIN SOURCES: PLANT-BASED Food Serving size Falafel 3.5 oz (100g), Hummus 2/3 cup (162 g) Soybeans, roasted (soy nuts) 1/3 cup (57g) Tempeh, cooked 3.5 oz (100 g) Tofu, firm 1 cup (252 g) kcal 333 Prot (g) 13.3 Carb (g) 31.8 Fat (g) 17.8 Leu (g) 0.9 269 12.8 23.2 15.6 0.9 256 22.4 18.5 12.2 1.8 196 18.2 9.4 11.4 1.4 176 20.6 4.2 10.6 1.8 Carb (g) 18.0 Fat (g) 20.5 Leu (g) 1.4 HIGH-FAT PROTEIN SOURCES: PLANT-BASED Food Serving size kcal Almonds 542 575 12.5 10.8 58.3 1.0 553 13.9 29.6 41.9 1.2 Chia seeds, whole 2/3 cup, whole (94 g) 2/3 cup, whole (88 g) 2/3 cup, whole (100g) 2/3 cup (100 g) Prot (g) 20.0 490 15.6 43.8 30.8 1.3 Flaxseed, whole 2/3 cup (111 g) 592 20.3 32.0 46.7 1.4 Hazelnuts 2/3 cup (76 g) 477 11.3 12.7 46.1 0.8 Peanuts 2/3 cup (96 g) 564 22.8 20.7 47.8 1.5 Pistachios 2/3 cup (81 g) 463 17.4 22.4 37.3 1.3 Pumpkin or squash seeds, unshelled Sunflower seed kernels, dry roasted 2/3 cup (91g) 493 22.4 16.2 41.8 1.9 2/3 cup (84 g) 492 16.3 20.3 42.1 1.2 Brazil nuts Cashews - 107 - Postscript Congrats, you made it! For those who finished the book and have questions, suggestions, criticisms, or compliments, feel free to email me: alaneats@gmail.com. Your feedback is muchappreciated, and will be helpful for improving future editions. This book is a free gift for current AARR subscribers. For those who are not yet subbed, but want to stay on top of the latest research (with instant access to the archive of monthly content dating back to 2008), go here: https://alanaragon.com/aarr Thanks & all the best, – Alan - 108 - References 1. 2. 3. 4. 5. 6. Wolfe RR, Baum JI, Starck C, Moughan PJ. Factors contributing to the selection of dietary protein food sources. Clin Nutr. 2018 Feb;37(1):130-138. [PubMed] Berryman CE, Lieberman HR, Fulgoni VL 3rd, Pasiakos SM. Protein intake trends and conformity with the Dietary Reference Intakes in the United States: analysis of the National Health and Nutrition Examination Survey, 20012014. 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[PubMed] ___________________________________________________________________________________________________ This is the end of the book, but the beginning of an adventure. - 142 -