Past, Present, and Future of Protein Nutrition of Dairy Cattle

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1 Past, Present, and Future of Protein Nutrition of Dairy Cattle Chuck Schwab Schwab Consulting, LLC, Boscobel, WI Professor Emeritus, Animal Sciences University of New Hampshire

2 Greetings from Wisconsin!

3 Past

4 Change has come slowly

5 A lot was known about protein nutrition 100 years ago [Feeds and Feeding (Henry and Morrison, 1917), J. Dairy Sci. ( ) For example, it was recognized at that time that: CP was a proximate nutrient Proteins differed in size, shape, solubility, function, digestibility and amino acid (AA) composition Dietary protein provided AA to the animal and feed proteins differed in nutritional value because of differences in AA composition Enzymes were needed for protein digestion and that AA had to be released before intestinal absorption could occur Absorbed AA were carried via the blood to tissues for protein synthesis Absorbed AA could be metabolized to glucose and, via glucose, could be used for energy and fat synthesis

6 Continued Feed NPN was composed of many different small compounds such as peptides, free AA, nucleic acids, amides, amines, and ammonia Grasses and legume forages contained the highest and most variable concentrations of NPN and that hays, and especially silages, contained more NPN than the same feed when fresh Many feeding standards emerged Savage A study of the feeding standards for milk production. Bul.Cornell, NY Agr. Expt. Sta Eckles Nutrients required for milk production. Res. Bul. Mo. Agr. Expt. Sta. 7. Haecker Feeding dairy cows. Bul. Minn. Agr. Expt. Sta Armsby The nutrition of farm animals. McCandish, A modification of the Haecker and Savage Feeding Standards for dairy cattle. dcp, TDN (dcp + dcho times dfat) dtrue protein, NE dcp, dcho, dfat dtrue protein, NE dcp, dcho

7 Emergence of a simplified feed table McCandish, 1920

8 Schwab and Broderick (2017)

9 Classification of amino acids Essential (EAA) 1. Arginine 2. Histidine 3. Isoleucine 4. Leucine 5. Lysine 6. Methionine 7. Phenylalanine 8. Threonine 9. Tryptophan 10. Valine Non-essential (NEAA) 1. Alanine 2. Aspartic acid 3. Asparagine 4. Cysteine 5. Glutamic acid 6. Glutamine 7. Glycine 8. Proline 9. Serine 10. Tyrosine Black et al., 1952

10 Schwab and Broderick (2017)

Evolution of studies measuring contribution of rumen bacteria to total AA supply of dairy cows This effort was led by the findings of Virtanen (1966), a Nobel Prize Laureate, who reported that all of

11 Evolution of studies measuring contribution of rumen bacteria to total AA supply of dairy cows This effort was led by the findings of Virtanen (1966), a Nobel Prize Laureate, who reported that all of the AA in milk protein were enriched with N 15 when the rumen was dosed with ( 15 NH 4 ) 2 SO 4 and that cows could reproduce and produce milk when all dietary CP was supplied by urea and ammonium salts This work stimulated efforts to measure ruminal outflows of microbial protein and rumen escape protein (i.e., RUP) in the abomasum or proximal duodenum

12 Evolution of studies measuring contribution of rumen bacteria and RUP to total AA supply of dairy cows Fraught with initial challenges, results from the first experiments using dairy and beef cattle began appearing in the early 1980 s An omasal sampling technique was introduced in the late 1990 s Information from these experiments has been used for building nutritional models (e.g., NRC, 2001), evaluating their predictive ability (e.g., Pacheco et al., 2012), and for deriving new prediction equations (e.g., White et al., 2017). The updated data set used by White et al. (2017) contains 550 treatment means from 147 studies using dairy cows

13 Effects of Rumen-Undegradable Protein on Dairy Cow Performance: A 12-Year Literature Review The data strongly suggest that increased RUP per se in dairy cow diets, which often results in a decrease in RDP and a change in absorbed AA profiles, does not consistently improve lactational performance Santos et al. (1998)

appearing in the early 1980 s An omasal sampling technique was introduced in the late 1990 s Information

g., Pacheco et al., 2012), and for deriving new prediction equations (e.g., White et al., 2017).

14 Evolution of studies measuring contribution of rumen bacteria and RUP to total AA supply of dairy cows Fraught with initial challenges, results from the first experiments using dairy and beef cattle began appearing in the early 1980 s An omasal sampling technique was introduced in the late 1990 s Information from these experiments has been used for building nutritional models (e.g., NRC, 2001), evaluating their predictive ability (e.g., Pacheco et al., 2012), and for deriving new prediction equations (e.g., White et al., 2017). The updated data set used by White et al. (2017) contains 550 treatment means from 147 studies using dairy cows

15 Schwab and Broderick (2017)

such as type of N supplements and carbohydrate fermentability Ammonia concentrations required to maximize ruminal OM digestion are at least as high as those required to maximize

16 Evolution of studies showing the importance of maintaining adequate rumen ammonia levels Schwab et al. (2005) reviewed the literature and concluded: Required ruminal ammonia concentrations can vary from 5 to 11 mm The optimum concentration is diet dependent and influenced by factors such as type of N supplements and carbohydrate fermentability Ammonia concentrations required to maximize ruminal OM digestion are at least as high as those required to maximize ruminal synthesis of microbial protein Not only is average ammonia concentration important, but also the time the concentration remains below some critical level Higher ruminal ammonia concentrations may be needed if more readily fermentable carbohydrates are fed

17 Schwab and Broderick (2017)

18 Trial Evolution of studies looking at limiting AA for lactating dairy cows of AA (Schwab et al., 1976) Arg His EAA composition of infusates (not including negative or positive controls) Thr Phe Leu Leu His Phe Ile Ile Ile Ile Ile Thr Val Val Val Val Val Val Met Met Met Met Met Met Met Met Met Lys Lys Lys Lys Lys Lys Lys Lys Lys % of response obtained with positive control infusions (all EAA or casein) Ave

19 Schwab and Broderick (2017)

20 Schwab and Broderick (2017)

21 Present

Rumen microorganisms have AA and ammonia requirements and cows have AA requirements 1) RDP (mixture of true protein and NPN) purpose is to meet the AA and ammonia requirements of rumen

22 Rumen microorganisms have AA and ammonia requirements and cows have AA requirements 1) RDP (mixture of true protein and NPN) purpose is to meet the AA and ammonia requirements of rumen microbes for maximum carbohydrate digestion and synthesis of microbial protein 2) RUP purpose is to provide the additional AA that the cow requires that are not provided by microbial protein

23 Factors affecting RDP requirements 1) Dietary content, quality, and mixture of fermentable carbohydrates Fermentable carbohydrates RDP Microbial protein VFA s 2) Quality of RDP (proportional supplies of true protein, short peptides, free AA and ammonia)

24 Factors affecting RUP requirements 1) Microbial protein yield 2) RUP digestibility 3) RUP-Lys digestibility (usually lower than RUP digestibility) 4) Amino acid composition of RUP Effect of levels of Lys and Met in MP on amounts of MP and RUP required to provide 180 g MP-Lys and 60 g MP-Met Lys/Met in MP (%) MP required (g/d) Required MP from RUP (g/d) Required RUP (g/d) Required RUP (% DM) 5.7/ / / /

25 Lys, Met and His concentrations in milk, rumen microbes and feedstuffs (% of CP), relative to estimated ideal concentrations in MP Lys Met His Lys Met His Milk Brewer s grains Bacteria Canola meal Protozoa Corn DDGS Corn gluten feed Ideal (Lapierre) Corn gluten meal Cotton seed Alfalfa silage Linseed meal Corn silage Soybean meal Grass silage Blood meal Barley Feather meal Corn Fish meal Wheat Meat meal

26 Response to lysine Protein yield response, g/d LysDI, % PDI From 557 cows Rulquin et al. (1993)

27 Optimum content of Lys in MP Milk protein content responses, g/100 g Percent Lys in MP (Met > 1.95 of MP)

28 Response to methionine Protein yield response, g/d MetDI, % PDI From 265 cows Rulquin et al. (1993)

29 Optimum content of Met in MP 0.20 Milk protein content responses, (g/100 g) Percent Met in MP (Lys > 6.50 of MP)

30 Required concentrations of Lys and Met in MP for maximal content of milk protein Model Lys Met Optimal Lys/Met ratio NRC (2001) Original release Revised v CPM-Dairy CNCPS Prior v v v Schwab et al. (2009), 2 Whitehouse et al. (2009), 3 Whitehouse et al. (2013), 4 Van Amburgh et al. (2015)

Amino acid balancing Definition A deliberate attempt, through selective use of protein supplements and RP-AA supplements, to achieve an amount

31 Amino acid balancing Definition A deliberate attempt, through selective use of protein supplements and RP-AA supplements, to achieve an amount and profile of absorbed AA that comes as close as possible to meeting the cows requirements for optimal health and performance without wasting AA

32 Transition cows mobilize body protein Metabolizable Protein Individual values were calculated from daily measurements of CP intake and milk yield and weekly milk composition of 80 cows fed a 17.8% CP ration (1.7 Mcal/kg NEL). Bell et al. (2000)

33 Abomasal infusion of casein into postpartum transition cows increases milk yield 1,2 Item 4 DIM 15 DIM 29 DIM DM intake MP, g/d Milk, kg/d ** ** ** ECM, kg/d ** * Protein, % Fat, % Urea, mm ** ** * BW change, kg ** P < 0.01, *P < 0.05 Larson et al. (2014)

34 Ruminant Lys and Met supplements Lys supplements AjiPro -L AminoShure-L Bovi-Lysine Lysine 35 LysiPEARL Megamine-L MetaboLys NoviLys USA Lysine Met supplements Smartamine M Mepron AminoShure-M MetaboMet MethioPlus MetiPEARL Novimet EB-Met MetaSmart (HMBi) Alimet Rhodimet AT 88 MFP (CaMHA) RP-Met Met analogs

35 Amino acid balancing is being rapidly adopted in the US 2,000 Cows fed AA balanced rations (000s) North & Central America 1, ,600 1, , , Adisseo, 2016

36 Benefits of AA balancing 1. Reduced need for MP and therefore, dietary RUP

37 Effect of amounts of Lys and Met in MP on amounts of MP and RUP required to provide 180 g of MP-Lys and 60 g of MP-Met Lys/Met in MP (%) MP required (g/d) Required MP from RUP (g/d) Required RUP (g/d) Required RUP (% DM) 5.7/ / / / ) Assumed microbial protein supply will not change when RUP supply is reduced, which is not correct 2) A benefit of reduced RUP feeding is the opportunity to feed more of the rest of the diet which will increase microbial protein synthesis. Therefore, the required RUP values are lower than indicated for last 3 diets 3) AA balancing is the only option for increasing the efficiency of use of RUP

38 Benefits of AA balancing 1. Reduced need for MP and therefore, dietary RUP 2. Reduced risk of a deficiency of the most limiting AA (e.g., Met and Lys) a. Increased milk yield, particularly in early lactation cows (2 to 4.0 kg/d more are common) b. Increased milk components (0.10 to 0.20% unit increases in protein and 0.10 to 0.15% unit increases in fat are common) c. Healthier and more productive transition cows

Better performance with Met supplementation during the transition period 1 Control MetaSmart 2 Smartamine M 2 DMI, kg/d 13.3 15.2 15.6 Milk, kg/d 35.7 b 38.1 ab 40.

39 Better performance with Met supplementation during the transition period 1 Control MetaSmart 2 Smartamine M 2 DMI, kg/d Milk, kg/d 35.7 b 38.1 ab 40.0 a Milk protein, % 3.04 b 3.26 a 3.19 ab Milk fat, % ECM, kg/d 41.0 b 44.8 a 45.0 a 1 Osorio et al. (2013) 2 Fed in amounts to achieve a predicted Lys/Met ratio in MP of 2.80/1

40 Liver lipids and whole-blood leukocyte phagocytosis Diet P -value CON MS SM Met 1 Liver, % wet wt Total lipid TAG Phagocytosis 2, % Contrast statement of CON versus MS + SM 2 Percent of immune cells able to engulf pathogens Osorio et al. (2014)

41 Biomarkers of liver function, inflammation and oxidative stress Diet P -value CON MS SM Diet Met 1 Liver function Carnitine, nmol/g tissue <0.01 Albumin, g/l Inflammation Ceruloplasmin, umol/l SAA, ug/ml Oxidative stress ORAC, mol/l Glutathione, mm Contrast statement of CON versus MS + SM Osorio et al. (2014)

42 Effect of improved methionine nutrition on reproduction Table 1. Amniotic vesicle size n Volume (mm 3 ) Primiparous Control RPM P value 0.71 Multiparous Control RPM P value 0.05 Toledo et al. unpublished

43 Effect of improved methionine nutrition on reproduction Table 2. Embryo size Primiparous n Crown-rump length (mm) Abdominal diameter (mm) Volume (mm 3 ) Control RPM P value Multiparous Control RPM P value Toledo et al. unpublished

44 Effect of improved methionine nutrition on reproduction Table 3. Pregnancy loss Primiparous Multiparous Interval Control RPM P Control RPM P d 12.8% 14.6% % 6.1% 0.03 (5/39) (6/41) (10/51) (3/49) Toledo et al. unpublished

45 Effect of maternal methionine supplementation on embryo quality and gene expression 76 genes out of 10,662 were up-regulated 200 genes were down-regulated Penagaricano et al. (2013)

47 Future

48 Future of protein nutrition 1) Feeding less rumen escape protein and more rumen protected AA supplements. 2) A greater availability of rumen protected AA supplements and better accountability of product marketing

49 Ruminant Lys and Met supplements Lys supplements AjiPro -L AminoShure-L Bovi-Lysine Lysine 35 LysiPEARL Megamine-L MetaboLys NoviLys USA Lysine Met supplements Smartamine M Mepron AminoShure-M MetaboMet MethioPlus MetiPEARL Novimet EB-Met MetaSmart (HMBi) Alimet Rhodimet AT 88 MFP (CaMHA) RP-Met Met analogs

50 Take home messages 1. DO NOT feed a rumen protected AA supplement unless you have seen confirmed estimates of bioavailability obtained with lactating cows using the plasma AA dose-response method 2. CONFIRM that the RPAA supplement was mixed and consumed with the rest of the diet as part of its evaluation 3. DO NOT accept milk production related data as proof of efficacy of a product such data favors inferior products when comparisons are made with superior products

University of New Hampshire approach for using the plasma free AA dose-response approach High producing cows are fitted with ruminal cannula for abomasal infusion of the unprotected AA Latin square

51 University of New Hampshire approach for using the plasma free AA dose-response approach High producing cows are fitted with ruminal cannula for abomasal infusion of the unprotected AA Latin square experiments with a minimum of 7-d periods (4-d adjustment, last 3 d for blood sampling) RPAA supplements are mixed in the TMR 8 h before feeding Cows are fed at 8-h intervals and milked at 12-h intervals Blood samples are collected last 3 days of each period at 2, 4, 6 and 8 h after morning feeding. Deproteinized plasma samples are pooled within day (across 4 sampling times) for each cow before AA analysis

52 Total Sulfur Amino Acids, µm Changes in plasma free sulfur AA concentrations with increasing amounts of infused or fed Met Infusion Smartamine M y = 1.98x R² = y = 1.64x R² = Bioavailability: 1.64/1.98 = x 100 = 82.8% Methionine infused or fed, g/d 1 Two 5 x 5 replicates (2013, 2014) Chirgwin et al. (2015)

53 A comparison of the relative bioavailabilities of AjiPro-L 2G and 3G Values for 2G and 3G were 37.9 and 42.4%, respectively. Slopes were not significantly different. Whitehouse et al. (2017)

54 Changes in plasma sulfur AA and HMTBa concentrations with increasing amounts of infused HMTBa Whitehouse et al. (2018)

55 Point is If you want X more grams of metabolizable Lys, you need to know how much of the Lys supplement you need to feed to get it If you want X more grams of metabolizable Met, you need to know how much of the Met supplement you need to feed to get it

56 Future of protein nutrition 1) Feeding less rumen escape protein and more rumen protected AA supplements. 2) A greater availability of rumen protected AA supplements and better accountability of product marketing 3) Better prediction of microbial protein synthesis and ruminal escape protein 4) Better definition of the intestinal availability of AA from microbial protein and rumen escape protein

57 Lactating dairy cattle in North America typically capture 25% of consumed dietary N in milk and tissue (Hristov et al., 2004). Accurate predictions of the intestinal supply of AA from dietary ingredients escaping ruminal degradation and from microbial protein are necessary to more precisely match the dietary supply of individual AA to requirements. Improving models used to balance dairy rations for AA will optimize milk protein yield and increase N efficiency (Khezri et al., 2011). Several in vitro, in situ, and in vivo methods of assessing protein and AA supplies to the animal have been developed; however, they all have limitations. This method appears to provide an accurate and precise in vivo assessment of individual AA plasma entry rates that can be used to better characterize individual feed ingredients in ruminants. Such information will result in more robust economic assessments of feeds and increased precision of diet formulation.

58 Future of protein nutrition 1) Feeding less rumen escape protein and more rumen protected AA supplements. 2) A greater availability of rumen protected AA supplements and better accountability of product marketing 3) Better prediction of microbial protein synthesis and ruminal escape protein 4) Better definition of the intestinal availability of AA from microbial protein and rumen escape protein 5) Better definition of ideal profile of absorbed AA and AA requirements

What is the ideal balance of absorbed AA for lactating cows? Amino acid Rulquin (2001) Doepel et al. (2004) Segmented Logistic linear model model Rulquin et al. (2007) Arg 3.1 4.8 4.6 4.5 His 3.0 2.

59 What is the ideal balance of absorbed AA for lactating cows? Amino acid Rulquin (2001) Doepel et al. (2004) Segmented Logistic linear model model Rulquin et al. (2007) Arg His Ile Leu Lys Met Phe Thr Trp Val Values obtained by integrating data from experiments in which individual and combinations of AA were infused postruminally in lactating cows

60 More recent thoughts on the ideal profile of EAA in MP 30 kg milk, 3.00% protein 60 kg milk, 3.00% protein Increase in MP Amino acid g/d % of MP g/d % of MP % Arginine Histidine Isoleucine Leucine Lysine Methionine Phenylalanine Threonine Tryptophan Valine Lapierre (2016)

61 OPTIMUM SUPPLY OF EACH EAA RELATIVE TO ENERGY Ryan Higgs, 2014

62 OPTIMUM SUPPLY OF EACH EAA RELATIVE TO ENERGY RULQUIN et al. (1993): 14.7 and 5.3% of EAA SCHWAB (1996): 14.9 and 5.1% of EAA Ryan Higgs, 2014

63 Future of protein nutrition 1) Feeding less rumen escape protein and more rumen protected AA supplements. 2) A greater availability of rumen protected AA supplements and better accountability of product marketing 3) Better prediction of microbial protein synthesis and ruminal escape protein 4) Better definition of the intestinal availability of AA from microbial protein and rumen escape protein 5) Better definition of AA requirements 6) Driven by better nutritional models and non-linear optimizers, better farm management practices, and issues of farm profitability and agricultural sustainability

64 Chuck Schwab Schwab Consulting, Boscobel, Wisconsin USA Professor Emeritus, Animal Sciences University of New Hampshire

Advances in Protein and Amino Acid Nutrition: Implications on Transition Cow Performance

Advances in Protein and Amino Acid Nutrition: Implications on Transition Cow Performance Chuck Schwab Schwab Consulting, LLC, Boscobel, WI Professor Emeritus, Animal Sciences University of New Hampshire