Substituting ground woody plants for cottonseed hulls in lamb feedlot diets:

Growth performance, blood serum chemistry, and rumen fluid parameters1

T. R. Whitney,*2 J. L. Glasscock,* J. P. Muir,† W. C. Stewart,‡ and E. J. Scholljegerdes§
*Texas A&M AgriLife Research, 7887 U.S. Hwy 87 N, San Angelo 76901; †Texas A&M AgriLife Research, 1229 N.
U.S. Hwy 281, Stephenville 76401; ‡Department of Animal and Range Sciences, Montana State University, Bozeman
59717; and §Animal and Range Sciences Department, New Mexico State University, Box 30003, Las Cruces 88003

ABSTRACT: Effects of using ground woody plants in

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Rambouillet wether lamb feedlot diets on growth per-
formance, blood serum, and rumen parameters were
evaluated. A randomized design study was used with
2 feeding periods (70% concentrate diet from d 0 to 27
[Period 1] and 86% concentrate diet from d 28 to 57
[Period 2]); the concentrate portion of the diets mainly
consisted of dried distiller’s grains with solubles and
sorghum grain. Lambs were individually fed 6 diets
that differed only by roughage source (n = 8 animals/
treatment; 32.9 ± 3.2 kg initial BW): either cottonseed
hulls (CSH; control) or ground wood consisting of
redberry juniper (RED), blueberry juniper (BLUE),
one-seeded juniper (ONE), or eastern red cedar (ERC)
Juniperus spp. or honey mesquite (MESQ; Prosopis
glandulosa). Using ground wood vs. CSH as the
roughage source did not affect (P > 0.12) BW. There
tended to be a treatment × day interaction (P = 0.07)
for lamb DMI, attributed to Period 1 when DMI was
greater (P < 0.05) for lambs fed CSH vs. RED, ONE,
ERC, or MESQ during the first 14 d and greater (P <
0.05) during d 14 to 28 vs. lambs fed ERC. Overall,
Key words: feedlot diet, juniper, lambs, mesquite, plant secondary metabolites, serum chemistry
ADG was less for lambs fed ERC (P < 0.10) or MESQ
(P < 0.05) vs. lambs fed CSH, but G:F was similar
(P > 0.10) among all lambs. Dietary treatments did
not affect (P > 0.15) ruminal pH, but treatment × day
interactions (P < 0.05) were observed for rumen fluid
ammonia N or molar proportions of propionate and
butyrate; few differences were observed within day.
However, overall, lambs fed RED or MESQ had great-
er (P < 0.05) total rumen VFA than lambs fed CSH. A
treatment × day interaction (P = 0.04) was observed
for the acetate:propionate ratio, but no differences
(P > 0.10) were observed within day. Treatment × day
interactions (P < 0.05) were observed for blood serum
glucose, γ-glutamyl transferase, the albumin:globulin
ratio, total bilirubin, β-hydroxybutyrate, P, Cl, and Mg,
with most results being less for lambs fed the wood-
based diets than for lambs fed the CSH diets. Results
suggested that even though lamb DMI was reduced
during the growing period when diets contained
30% ground woody products (RED, ONE, ERC, and
MESQ), animal health and rumen fluid parameters
were not negatively affected by ground woody plants.

© 2017 American Society of Animal Science. All rights reserved. J. Anim. Sci. 2017.95:4150–4163
doi:10.2527/jas2017.1649
INTRODUCTION

Feed price fluctuation and seasonality of traditional

1This work was supported by the USDA National Institute
of Food and Agriculture Hatch Project number 205866 and was
funded in part by the National Sheep Industry Improvement Center
(NSIIC). Appreciation is expressed to POET Nutrition (Sioux Falls,
SD) for donating dried distiller’s grains with solubles.
2Corresponding author: trwhitney@ag.tamu.edu
Received April 19, 2017.
Accepted July 21, 2017.
4150
feed ingredients are major concerns of livestock feed-
ers. The price of dietary roughage components such
as hay and cottonseed hulls can dramatically fluctu-
ate within any given year and, at times, be completely
unavailable in certain regions. This is especially true
during drought and times when greater quantities of
roughages are purchased by nonagriculture industries.
However, ground woody products such as Juniperus
spp. and Prosopis spp. are not subject to seasonal feed
 Ground woody products as roughage sources
pricing and availability, require no inputs by man to es-
tablish and grow, and are the only feed ingredients that
can increase rangeland forage production after being
harvested (Carter, 1958; Coultrap et al., 2008). Use of
ground wood as roughage in livestock feed is not a new
concept (Maynard, 1920; Marion et al., 1957). More re-
cently, ground Juniperus pinchotii and Juniperus ashei
have been effectively used in lamb (Whitney et al., 2014)
and goat (Glasscock et al., 2015) feedlot diets and sup-
plements (Stewart et al., 2017) and have enhanced meat
sensory characteristics (Whitney and Smith, 2015).
Millions of hectares of North American range-
lands are infested with Juniperus spp. and Prosopis spp.
plants (Felker, 1996; Ansley et al., 2006). Woody plant
encroachment can negatively affect livestock produc-
tion and natural resources (Scholes and Archer, 1997);
however, converting these plants into feed ingredients
would help offset removal costs while synergistically
reducing livestock feed costs. Economic incentives will
be created for multiple industries if the feeding value
of ground woody products is found to be comparable
to similar ingredients. Therefore, the objective was to
evaluate the use of ground woody plants in mixed diets
on lamb growth performance, health, and rumen param-
eters. The hypothesis was that woody plants would be
an effective, safe, and economically viable alternative
to cottonseed hulls in lamb feedlot production systems.
MATERIALS AND METHODS
Animals and Management
The experimental protocol was approved by the
Texas A&M AgriLife Research Agriculture Animal
Care and Use Committee (number 2014-006); College
Station, TX). Forty-eight Rambouillet wether lambs
(approximate age of 4 mo and 32.9 ± 3.2 kg initial
BW) previously had been on pasture; therefore, a 19-d
feed yard adaptation period was used. Unshorn lambs
received an ear tag, subcutaneous clostridial vaccine
(Vision 7 with SPUR; Intervet Inc., Omaha, NE), and
an oral dose of levamisole (Prohibit; AgriLabs, St.
Joseph, MO). During the first 12 d of the adaptation
period, lambs were group-fed and had ad libitum ac-
cess to oat hay, which was supplemented with a 60%
concentrate diet (0.22 kg/(d·lamb); DM basis).
Each lamb was weighed 7 d before study initiation,
stratified by BW, and randomly assigned to an individu-
al, completely covered dirt pen (2.44 by 2.97 m) with an
automatic watering system and a feed bunk. Each lamb
was also randomly assigned to a treatment diet (n = 8
lambs/treatment) that differed only by roughage source:
either cottonseed hulls (CSH) or a ground woody prod-
uct consisting of J. pinchotii (redberry juniper [RED]),
4151
J. ashei (blueberry juniper [BLUE]), Juniperus mono-
sperma (one-seeded juniper [ONE]), Juniperus virgin-
iana (eastern red cedar [ERC]), or Prosopis glandulosa
(honey mesquite [MESQ]). During the last 7 d of the
adaptation period, lambs did not receive any hay but
were fed a common 66% concentrate diet that was grad-
ually replaced with the appropriate treatment diet.
During Period 1 (d 0 to 27), lambs were fed
their respective 70% concentrate treatment diet. All
mixed diets were nonagglomerated, contained 22 g
monensin/t of feed (Rumensin 90; Elanco Animal
Health, Indianapolis, IN), and were fed once daily at
0900 h with an approximate allowance of 10% refusal.
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Lambs were then transitioned over 4 d into Period 2 (d
28 to 57), onto their respective 86% concentrate treat-
ment diet, by gradually replacing the Period 1 diet with
the Period 2 diet. Lamb BW was recorded on d 0, 14,
28, 43, and 57; lamb BW was also recorded after lambs
were shorn on d 57. Average daily gain and average
daily DMI were determined between days that BW was
recorded, and G:F was calculated between weigh days
by dividing ADG by average daily DMI. Blood serum
was collected on d 0. Ruminal fluid and blood serum
was also collected on d 14 to 15 and d 52 to 53; 4 lambs/
treatment were sampled on consecutive days.
Cost per metric ton of feed (as-fed basis) was cal-
culated using ingredient prices based on local markets.
The price of juniper (US$80/dry t, freight on board)
was based on estimated processing costs, after consult-
ing with juniper processing specialists; transportation
costs were not included for any ingredient. For each
treatment group, average daily feed cost per kilogram
of BW gain was calculated on a DM basis as (DMI
needed to gain 1 kg of BW, kg × $/kg of feed).
Sample Collection and Measurements
Woody Plant Harvesting, and Feed Processing,
Collection, and Analysis. During May, the entire
aboveground biomass from mature Juniperus spp.
(including leaves) and mature P. glandulosa (exclud-
ing leaves) trees were harvested separately by species,
chipped (Model BC1800, Vermeer Corp., Pella, IA),
and dried for 5 h to approximately 93% DM using a
drying trailer equipped with a perforated metal bottom
sieve and a jet dryer (26 to 31°C; model 2001; Peerless
Manufacturing Co., Shellman, GA). Chipped material
was hammer milled to pass a 4.76-mm screen (Sentry,
model 100; Mix-Mill Feed Processing Systems, Bluffton,
IN), bagged, and stored under cover. Additional feed
subsamples were dried to constant weight in a forced-air
oven at 103°C to determine DM concentration.
Nutritive characteristics of woody plants were eval-
uated using random subsamples of mechanically dried
4152 Whitney et al.

Table 1. Chemical composition and digestibility (% DM basis) of cottonseed hulls (CSH), sorghum grain, and
dried distiller’s grains with solubles and ground Juniperus spp. and Prosopis glandulosa used in the treatment diets
Ingredient2
Item1 Cottonseed hulls Sorghum grain Corn DDGS J.pin J.ash J.mon J.vir P.glan
Nutrient composition
DM, % 92.3 92.6 91.8 93.9 93.8 95.4 93.8 91.9
CP, % 3.5 11.9 30.4 2.9 2.8 2.5 3.8 5.7
ADICP, % 3.2 1.5 1.3 1.5 1.6 1.4 1.8 2.5
aNDF, % 85.2 7.0 30.4 62.1 65.0 71.0 68.0 74.7
ADF, % 62.1 5.3 12.9 49.4 52.1 57.9 55.8 57.8
Lignin, % 16.4 0.9 2.9 19.4 21.2 23.2 21.7 17.9
Crude fat, % 0.6 3.1 8.7 3.2 3.2 4.5 4.1 6.2
Ash, % 3.6 3.6 4.7 4.9 4.8 3.4 4.4 4.3
Ca, % 0.12 0.04 0.07 1.31 1.53 1.27 1.37 1.42

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P, % 0.04 0.21 0.88 0.04 0.04 0.03 0.06 0.03
S, % 0.06 0.14 .93 0.07 0.04 0.04 0.05 0.06
K, % 0.99 0.34 1.33 0.33 0.16 0.11 0.24 0.35
Mg, % 0.14 0.12 0.38 0.06 0.04 0.03 0.1 0.02
Na, % 0.01 0.01 0.31 1.25 0.01 0.01 0.01 0.01
Fe, mg/kg 33 48 85 168 98 113 140 112
Zn, mg/kg 5 20 64 104 9 6 11 4
Cu, mg/kg 3 3 8 3 2 2 2 2
Particle distribution,3 %
19 to 8 mm 68.9 – – 0.20 3.9 3.7 5.2 0.3
8 to 1.18 mm 27.6 – – 69.3 68.1 68.2 63.7 70.3
<1.18 mm 3.5 – – 30.5 28.0 28.2 31.1 29.4
<8 mm 31.1 – – 99.8 96.2 96.3 94.8 99.7
CT, % 31.1 – –
Extractable 1.4 0 – 2.8 3.2 1.8 2.4 0.9
Protein bound 1.8 0 – 2.1 2.3 1.9 2.3 3.8
Fiber bound 0.2 0 – 0.0 0.0 0.0 0.0 0.1
Total 3.2 0 – 4.9 5.5 3.7 4.7 4.7
True IVDMD4 18.9 94.3 75.6 31.3 29.0 24.6 26.9 20.9
1ADICP = acid detergent insoluble CP; aNDF = neutral detergent fiber, using α-amylase and sodium sulfite; CT = condensed tannins.
2J.pin = Juniperus pinchotii; J.ash = Juniperus ashei; J.mon = Juniperus monosperma; J.vir = Juniperus virginiana (entire aboveground biomass);
P.glan = Prosopis glandulosa (entire aboveground biomass excluding leaves); DDGS = dried distiller’s grains with solubles (corn DDGS was produced
from corn ethanol production; POET, LLC, Sioux Falls, SD).
3Ingredients sieved using the manually operated Penn State Particle Separator (Lammers et al. (1996) according to modified procedures of Kononoff et al. (2003).
4True IVDMD represents true 48-h IVDMD.

and hammer milled (4.76-mm screen) woody plants.
Subsamples of CSH, sorghum grain, and dried dis-
tiller’s grains with solubles (DDGS) were collected 6
times throughout the trial; the first 3 and last 3 subsam-
ples were combined separately by period for analysis
(Table 1). Three random subsamples of treatment diets
(Table 2) were collected during both periods, combined
by period, and separately analyzed. These samples were
dried at 55°C in a forced-air oven (model 630; Napco,
Inc., South Haven, MI) for 48 h, ground through a 1-mm
screen (Wiley mill; Arthur H. Thomas Co., Philadelphia,
PA), and stored at −20°C. Nitrogen was analyzed us-
ing a standard method (method 990.03; AOAC, 2006),
and CP was calculated as 6.25 × N. The NDF and ADF
were analyzed according to procedures of Van Soest et
al. (1991), which were modified for an Ankom 2000
Fiber Analyzer (Ankom Technology Corp., Fairport,
NY) using α-amylase and sodium sulfite. In addition, N
was analyzed in residue remaining after the ADF proce-
dure and multiplied by 6.25 to determine acid detergent
insoluble CP. Standard methods were used to analyze
lignin (method 973.18; AOAC, 2006), crude fat (meth-
od 2003.05; AOAC, 2006), and ash (method 942.05;
AOAC, 2006). For individual mineral analyses, samples
were digesting with a Microwave Accelerated Reaction
System (MARS6; CEM Corp., Matthews, NC) and then
analyzed using a Thermo Jarrell Ash IRIS Advantage
HX Inductively Coupled Plasma Radial Spectrometer
(Thermo Instrument Systems Inc., Waltham, MA).
To evaluate particle size distribution of CSH and
the ground woody plants, as-fed subsamples of these
ingredients were sieved with the manually operated
 Ground woody products as roughage sources
Figure 1. Photograph represents material (not equal weights) that was
placed in warm water for 10 min after being stirred: (a) cottonseed hulls,
(b) ground Juniperus ashei (entire tree with leaves), (c) ground Prosopis
glandulosa (entire trees without leaves), and sieved cottonseed hulls with
particle distributions of (d) 19 to 8 mm, (e) 8 to 1.18 mm, or (f) <1.18 mm.
Penn State Particle Separator (Lammers et al., 1996)
according to modified procedures of Kononoff et al.
(2003). The percentage of material (as-fed) retained
on each sieve and under the 8-mm sieve was calcu-
lated based on total sieved material. In addition, to
visually demonstrate extreme differences in particle
buoyancy, nonsieved and sieved cottonseed material
(8 mm, 1.18 mm, and the bottom pan), along with the
J. ashei and P. glandulosa material, was placed into
warm water for 10 min and then photographed (Fig. 1).
Condensed tannins in the juniper, mesquite, CSH,
and sorghum grain were assayed for soluble, protein-
bound, and fiber-bound fractions using methods de-
scribed by Terrill et al. (1992); samples were oven-
dried, and standards were prepared for each individual
ingredient as recommended by Wolfe et al. (2008). An
Ankom model DaisyII incubator (Ankom Technology
Corp., Macedon, NY) was used to determine 48-h true
IVDMD by incubating individual ingredients and each
treatment diet in separate F57 bags (3 replicates; Ankom
Technology Corp., Macedon, NY) for 48 h. Each bag
contained 0.35 g of material that was ground to pass
a 2-mm screen (Wiley mill). The F57 bags of mixed
diets containing the ground woody material were incu-
4153
bated in separate jars from the cottonseed hull–based
(control) diet to eliminate any potential confounding ef-
fects. Bags were placed into jars containing 400 mL of
sheep rumen fluid (collected orally from 6 donors and
combined) and 1,600 mL of McDougall’s buffer solu-
tion (1.0 g of urea/L; McDougall, 1948). The sheep
donors were grazing native pasture common to the
Edwards Plateau area of Texas (e.g., Nassella leucotri-
cha, Hilaria belangeri, and Bouteloua spp.). One blank
bag per jar was also included and used to adjust for po-
tential residue on the bags. After anaerobic incubation
at 39°C, bags were gently rinsed under cold water for 5
min, subjected to the NDF procedure (using α-amylase
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and omitting sodium sulfite), gently rinsed in acetone,
dried at 55°C in a forced-air oven for 48 h, and weighed.
Rumen Fluid and Blood Serum Collection and
Analysis. A 10-mL blood sample was collected 4 h after
feeding from each lamb via jugular venipuncture using
a nonheparinized vacutainer collection tube (serum sep-
arator tube, gel, and clot activator; Becton, Dickinson
and Company, Franklin Lakes, NJ). Blood was allowed
to clot and then centrifuged (Beckman Coulter TJ6 re-
frigerated centrifuge; Beckman Coulter, Inc., Brea, CA)
at 970 × g for 25 min at 4°C. Serum was decanted and
frozen at −20°C until analyzed. Serum chemistry was
analyzed by The Texas A&M Veterinary Diagnostic
Laboratory, Amarillo, using an Olympus AU400E ana-
lyzer (Olympus America Inc., Center Valley, PA).
Rumen fluid was collected orally from each lamb
4 h after feeding. The pH of each subsample was im-
mediately recorded, and the remaining fluid was filtered
through 4 layers of cheesecloth into prewarmed ther-
moses purged with CO2. A subsample of the filtered flu-
id was immediately placed on ice and stored at −80°C.
Additional subsamples (1 mL) were acidified with 4 mL
of 0.1 N HCl (Farmer et al., 2004) and stored at −80°C
for analysis of ammonia N using a Beckman Coulter
DU640 spectrophotometer (Beckman Instruments, Inc.,
Fullerton, CA; methods of Broderick and Kang [1980])
and VFA using an Agilent 6890N gas chromatograph
(Agilent Technologies, Inc., Wilmington, DE; methods
of Baumgardt [1964] and Fritz and Schenk [1979]).
Statistical Analysis
Lamb BW, DMI, DMI as a percentage of BW,
ADG, and G:F were analyzed using the PROC MIXED
procedure of SAS (SAS Inst. Inc., Cary, NC) with a
model that included treatment, day, and treatment ×
day interaction; day was a repeated measure, individual
lamb was the subject, and Kenward–Roger approxima-
tion was used (and in all subsequent analysis). Suitable
covariance structures (unstructured [UN], compound
symmetry, compound symmetry heterogeneity) were
4154 Whitney et al.

Table 2. Ingredient, chemical composition (% DM basis), and digestibility of treatment diets

Diet2
Period 1 Period 2
Item1 CSH RED BLUE ONE ERC MESQ CSH RED BLUE ONE ERC MESQ
Cottonseed hulls 30.0 – – – – – 14.0 – – – – –
Ground wood – 30.0 30.0 30.0 30.0 30.0 – 14.0 14.0 14.0 14.0 14.0
Corn DDGS 40.0 40.0 40.0 40.0 40.0 40.0 40.0 40.0 40.0 40.0 40.0 40.0
Ground sorghum grain 21.7 21.7 21.7 21.7 21.7 21.7 37.6 37.6 37.6 37.6 37.6 37.6
Molasses, cane 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0
Limestone 2.2 2.2 2.2 2.2 2.2 2.2 2.3 2.3 2.3 2.3 2.3 2.3
Ammonium chloride 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5
Salt 0.6 0.6 0.6 0.6 0.6 0.6 0.6 0.6 0.6 0.6 0.6 0.6
Mineral and vitamin premix3 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0
Nutrient composition, %

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DM 91.9 90.8 90.6 91.6 91.4 91.2 90.6 89.6 89.9 90.1 90.0 89.7
CP 18.2 18.5 18.6 17.3 18.5 18.8 19.1 19.8 19.1 18.5 19.3 19.7
aNDF 32.6 34.8 33.2 37.9 34.6 36.6 25.4 25.7 23.1 26.3 25.6 26.6
ADF 16.4 19.9 19.0 22.7 20.4 19.7 13.5 13.6 12.9 14.7 13.3 12.1
Ca 1.2 1.4 1.4 1.3 1.3 1.4 1.2 1.4 1.3 1.3 1.2 1.4
P 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5
Ca:P ratio 2.3 2.7 2.9 2.8 2.9 3.0 2.4 2.7 2.8 2.7 2.5 2.9
Ash 8.1 8.0 8.5 7.7 8.5 8.4 7.8 9.1 8.3 8.0 8.1 8.5
True IVDMD4 70.8 64.4 67.1 60.8 65.0 63.6 74.6 72.6 74.7 74.8 73.2 73.8
1DDGS = dried distiller’s grains with solubles (corn DDGS was produced from corn ethanol production; POET, LLC, Sioux Falls, SD); aNDF = neutral
detergent fiber, using α-amylase and sodium sulfite..
2During Period 1 (d 0 to 27), lambs were fed a 70% concentrate diet. Lambs were transitioned over 4 d into Period 2 (d 28 to 57) onto an 86% concen-
trate diet. Treatment diets were nonagglomerated and ingredient composition differed only by roughage source: either CSH or ground woody products
(redberry juniper [RED; Juniperus pinchotii], blueberry juniper [BLUE; Juniperus ashei], one-seeded juniper [ONE; Juniperus monosperma], eastern
red cedar [ERC; Juniperus virginiana], or honey mesquite [MESQ; Prosopis glandulosa]). Juniperus (entire aboveground biomass) and Prosopis (entire
aboveground biomass except for leaves) species were chipped, dried, and hammer milled to pass a 4.76-mm sieve. Monensin (Rumensin 90; Elanco Animal
Health, Indianapolis, IN) was added to each diet at 22 g/t of feed.
3Mineral premix included NaCl; KCl; S; MnO; ZnO; vitamins A, D, and E; CaCO ; cottonseed meal; cane molasses; and animal fat (0.11 to 0.16% Ca,
3
0% P, 24.6 to 26.5% Na, 0% Mg, 9.57% K, 0% Se, and 64 to 69% salt).
4True IVDMD represents true 48-h IVDMD.

compared for each model, and UN was determined
to be the most appropriate. Data are reported as least
squares means with greatest SE (and in all subsequent
analysis described below). Differences in least squares
means between lambs fed compound symmetry hetero-
geneity and lambs fed treatment diets were evaluated
using the DIFF option with a SIMULATE adjustment.
When a treatment × day interaction was observed (P
< 0.10), effects were evaluated within that interaction.
However, all growth performance data are presented by
day due to the uniqueness of each feeding period.
Rumen fluid and blood serum parameters were ana-
lyzed using PROC GLIMMIX (SAS Inst. Inc.) with a
model that included treatment, day, and treatment × day
interaction; day was the random effect and individual
lamb was the subject. When a variable did not have a
treatment × day interaction but had a treatment effect
(alanine aminotransferase [ALT], creatinine, and Na),
the analysis was rerun with d 0 removed so that aver-
ages across days would reflect actual treatment effects.
Suitable covariance structures (UN, compound symme-
try, and compound symmetry heterogeneity for serum
parameters and autoregressive-1 and autoregressive
heterogeneity-1 for rumen parameters) were compared
for each model; compound symmetry heterogeneity was
determined to be the most appropriate structure for se-
rum and autoregressive heterogeneity-1 for rumen pa-
rameters). Various serum parameters (β-hydroxybutyrate
[BHB], urea N, NEFA, and globulin) and rumen fluid pa-
rameters (molar proportions of acetate, propionate, and
butyrate) did not have normal distributions; therefore,
lognormal distributions were used and data were post-
processed to the original scale. Statistical significance
was declared at P ≤ 0.05 and a tendency at 0.5 < P ≤ 0.10.
RESULTS AND DISCUSSION
Chemical and Physical Composition and Digestibility
of Individual Ingredients and Treatment Diets
The chemical composition of the ground woody
plants (Table 1) and CSH reveals their low-quality
 Ground woody products as roughage sources
roughage characteristics (e.g., low CP and high struc-
tural plant fiber characteristics; Fahey et al., 1994);
Stewart et al. (2015) reported similar results for the
Juniperus spp. Particle size distribution data reveals that
the neither CSH or any of the ground woody material
contained any particles greater than 19 mm (Table 1).
However, 69% of the cottonseed hull particles were re-
tained on the 8-mm sieve, whereas less than 5.3% of any
woody material was retained on the same sieve. In addi-
tion, woody material had approximately 30% very fine
particles (<1.18 mm) vs. only 3.5% for CSH. Figure 1
shows a crude evaluation of particle buoyancy of CSH
(and 3 different sized particles) and the woody products.
Even though numerous factors affect functional specific
gravity (particle size, rate of hydration, and digestibil-
ity), differences between CSH and the woody plants are
evident. Significance of particle size distribution and
buoyancy are discussed in the proceeding section.
The maximum condensed tannin (CT) concentra-
tion in any of the woody plant material or CSH used
in the current study was 5.5%, resulting in a maxi-
mum CT concentration of 1.65% in any of the mixed
diets. Results from previous studies suggested that 2
to 4% CT (DM basis) is beneficial to animal perfor-
mance (Min et al., 2003, 2012; Solaiman et al., 2010),
although bioactivity of CT is also a function of molec-
ular weight and stereochemistry, which contribute to
many diverse chemical structures across plant species
(Naumann et al., 2013), and such diversity of CT likely
exists across roughage ingredients in the current study.
Total volatile terpene oil was not analyzed, but me-
chanically dried woody products, similar to what was
used in the current study, have been reported to contain
less than 1.1% terpene oil (DM basis; Stewart et al.,
2015). Furthermore, even though the CSH and woody
plant material did not contain significant amounts of
CP, the mixed diets contained excess CP due to the use
of DDGS (Table 2). Considering a price (freight on
board) of $85/t for CSH and $80/t for ground woody
products, the cost of all the diets were similar within
period: $176/t in Period 1 and $201/t in Period 2.
Animal Performance
Lamb growth performance data are presented
in Table 3. There tended to be a treatment × day in-
teraction (P = 0.07) for lamb DMI, which is mainly
attributed to the first 14 d on feed, when lambs fed
CSH had greater (P < 0.05) DMI than lambs fed RED,
ONE, ERC, or MESQ (diets contained 30% CSH or
30% ground wood, DM basis). After d 14, all lambs
had similar (P > 0.10) DMI, except for lambs fed
ERC, which continued to have less (P < 0.05) DMI
than lambs fed CSH from d 14 to 28. A day effect (P <
4155
0.001) was also observed for lamb DMI: d 14 vs. 28
(P < 0.001), d 28 vs. 43 (P < 0.04), and d 43 vs. 57
(P < 0.01). Results of DMI as a percentage of BW
were similar to daily DMI, except between d 14 to 28,
when lambs fed RED had less (P < 0.05) and lambs
fed MESQ tended to have less (P < 0.10) DMI as a
percent of BW than lambs fed CSH (Table 3).
Dietary intake regulation is multifaceted and in-
volves interactions of physical and chemical feed
characteristics, which affect animal feeding behavior
such as sensory and satiety characteristics, along with
physiological mechanisms of the animal and its rumen.
Physical characteristics of feed ingredients such as par-
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ticle size, fragility, buoyancy, and rate of hydration can
affect physical effectiveness of NDF (peNDF), which
can ultimately affect DMI and animal growth and feed
efficiency (Hall and Akinyode, 2000; Kononoff and
Heinrichs, 2003). Even though physical characteris-
tics were not extensively evaluated in this trial, many
of these characteristics have been reported for CSH
(Mertens, 2002) and a similar woody product (J. pin-
chotii leaves and small stems; Whitney et al., 2014).
Physical characteristics and NDF concentrations of
CSH suggest that they are dense (Defoor et al., 2002),
contain less peNDF than many other forage-based in-
gredients (Mertens, 2002), and do not readily form a
ruminal mat layer (Owens and Goetsch, 1988; Moore
et al., 1990; Varga and Harpster, 1995). However, CSH
are palatable, have greater roughage value than alfal-
fa hay (Guthrie et al., 1996; Defoor et al., 2002), can
enhance rumination (Kononoff and Heinrichs, 2003),
and are an effective source of fiber for cattle and sheep
(Hall and Akinyode, 2000; Whitney and Lupton, 2010;
Whitney and Muir, 2010). Using similar types of lambs
fed DDGS-based diets, Whitney and Lupton (2010) re-
ported that DMI linearly increased as the percentage
of CSH increased from 10 to 30% in the diet. Other
trials have reported greater DMI when CSH are used
than when forages are used in mixed diets (Hall and
Akinyode, 2000), likely due to CSH increasing liquid
and passage rate of the grain (Cole et al., 1976; Moore
et al., 1990; Akinyode et al., 1999).
Grinding low-quality forages can increase passage
rate and DMI (Mertens, 1997), and it has been suggested
that particles smaller than 1.18 mm rapidly exit the ru-
men (Poppi et al., 1985; Mertens, 1997). Therefore, con-
sidering that 28 to 31% of wood particles were smaller
than 1.18 mm whereas CSH had only 3.5% particles
smaller than 1.18 mm (Table 1), it would be plausible
that wood particles would exit the rumen more rap-
idly, resulting in greater DMI, as previously described
(Lammers et al., 1996). However, grinding can also re-
duce peNDF and, thus, chewing activity (Lammers et al.,
1996). Furthermore, the majority of Juniperus spp. and
4156 Whitney et al.

Table 3. Effects of replacing cottonseed hulls (CSH) with ground woody products on lamb growth performance1
Diet2 P-value4
Item CSH RED BLUE ONE ERC MESQ SEM3 T D T×D
DMI, kg/d 0.004 <0.001 0.07
d 14 1.64 1.11* 1.39 1.19* 1.07* 1.22* 0.08
d 28 1.90 1.41 1.69 1.47 1.27* 1.45 0.10
d 43 1.81 1.53 1.62 1.64 1.48 1.56 0.09
d 57 1.70 1.40 1.43 1.61 1.45 1.47 0.09
d 0 to 57 1.76 1.36* 1.53 1.48 1.32* 1.43* 0.08
DMI, % of BW 0.003 <0.001 <0.001
d 14 4.32 3.02* 3.78 3.26* 3.12* 3.33* 0.15
d 28 4.55 3.56* 4.16 3.68 3.40* 3.63† 0.19
d 43 3.97 3.54 3.68 3.76 3.69 3.70 0.17
d 57 3.39 3.00 3.01 3.35 3.30 3.18 0.14

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d 0 to 57 4.06 3.28 3.66 3.51 3.38 3.46 0.13
ADG, kg 0.03 <0.001 0.28
d 14 0.39 0.25 0.32 0.25 0.15 0.22 0.04
d 28 0.27 0.20 0.27 0.22 0.27 0.24 0.04
d 43 0.25 0.22 0.22 0.25 0.21 0.15 0.04
d 57 0.61 0.50 0.49 0.60 0.49 0.45 0.07
d 0 to 57 0.38 0.29 0.33 0.33 0.28† 0.27* 0.03
G:F, kg/kg 0.66 <0.001 0.37
d 14 0.24 0.22 0.23 0.21 0.13 0.19 0.03
d 28 0.14 0.14 0.16 0.15 0.21 0.17 0.03
d 43 0.15 0.15 0.14 0.15 0.15 0.10 0.03
d 57 0.36 0.35 0.34 0.38 0.32 0.30 0.04
d 0 to 57 0.22 0.21 0.22 0.22 0.20 0.19 0.02
1During Period 1 (d 0 to 27), lambs were fed a 70% concentrate diet. Lambs were transitioned over 4 d into Period 2 (d 28 to 57) onto an 86% concentrate
diet. Within row means with a different superscript than the control diet (CSH) differ (P < 0.05); treatments were compared only with CSH, not to each other.
2Treatment diets were nonagglomerated and ingredient composition differed only by roughage source: either CSH or ground woody products (redberry
juniper [RED; Juniperus pinchotii], blueberry juniper [BLUE; Juniperus ashei], one-seeded juniper [ONE; Juniperus monosperma], eastern red cedar
[ERC; Juniperus virginiana], or honey mesquite [MESQ; Prosopis glandulosa]). Juniperus (entire aboveground biomass) and Prosopis (entire aboveg-
round biomass except for leaves) species were chipped, dried, and hammer milled to pass a 4.76-mm sieve.
3SEM represents the greatest SEM.
4T = treatment effect; D = day effect.

Within row: †0.05 > P < 0.10; *P < 0.05.

P. glandulosa particles are dense (Fig. 1), and particle
density can be great enough to reduce passage rate as
particles descend to the ventral rumen (King and Moore,
1957; Welch, 1986). Therefore, when compared with
lambs fed CSH, greater woody particle density, along
with reduced 48-h true IVDMD of the entire diet (Table
2), may have increased ventral rumen fill, contributing
to the observed reduction in DMI.
Figure 1 also illustrates that diets containing the
ground woody products would provide almost no ru-
men mat or particle stratification, which would reduce
efficient rumen function (Bernard et al., 2000; Mertens,
2002). This hypothesis is supported by Whitney et al.
(2014), who reported that ground J. pinchotii particles
(leaves and small stems) incubated in a ruminal fluid–buf-
fer mixture for 10 min or 2, 6, and 24 h resulted in 32, 12,
20, and 16% buoyant particles, respectively. Furthermore,
small, dense woody particles probably reduced effective-
ness of regurgitation and the structural integrity and num-
ber of daily boluses formed (Jaster and Murphy, 1983;
Pond et al., 1987; Luginbuhl et al., 1989), which would
reduce mastication of the larger woody particles. Using
40% DDGS in the mixed diets would exacerbate these
conditions, because DDGS is dense, and even though it
has greater NDF than typical concentrate sources, it can
reduce chewing activity (Zhang et al., 2010).
When DMI was greater for lambs fed CSH during
Period 1, it was likely a result of a more functional ru-
men environment, which enhanced chewing, rumination,
and particulate passage rate while reducing fill and the
energy dilution effect of low-quality fiber. Supporting
this conclusion is the fact that during the first 14 d of
the trial, calculated TDN, DE, and ME intakes (data not
shown) were met for lambs fed BLUE, barely met for
lambs fed ONE or MESQ, and not met for lambs fed
RED or ERC. Physical factors were likely suppressing
the increase in DMI, which has been observed when
dietary energy intake is reduced (Bartle et al., 1994;
 Ground woody products as roughage sources
Figure 2. Effects of replacing cottonseed hulls (CSH; control diet)
with ground woody plants on lamb BW. Treatment diets differed only by
roughage source; either CSH or ground woody products (redberry juniper
[RED; Juniperus pinchotii], blueberry juniper [BLUE; Juniperus ashei],
one-seeded juniper [ONE; Juniperus monosperma], eastern red cedar
[ERC; Juniperus virginiana], or honey mesquite [MESQ; Prosopis glan-
dulosa]). During Period 1 (d 0 to 27), lambs were fed a 70% concentrate
diet. Lambs were transitioned over 4 d into Period 2 (d 28 to 57) and fed an
86% concentrate diet. Data presented as least squares means ± SEM. No
treatment × day interaction (P = 0.71) or treatment (P = 0.12) effect was
observed, but a day effect (P < 0.001) was detected.
Forbes, 2003; Allen et al., 2009). Using less DDGS and
more of a concentrate ingredient that contains greater
starch may enhance effectiveness of the 30% woody-
based diets fed during Period 1. However, effects on
DMI due to CT in the CSH and woody plants and ter-
penes in the Juniperus spp. should not be discounted.
Condensed tannin concentration within diets was
likely not a significant contributing factor in reducing
DMI considering that 1) DMI was similar (P > 0.10)
among all lambs during the preconditioning period as the
quantity of each treatment diet was increased (data not
shown), 2) plants containing much greater CT concentra-
tions than the current trial either enhanced (goats; Terrill
et al., 2007; Moore et al., 2008) or did not negatively
affect DMI or growth performance (lambs; Chafton,
2006), 3) lambs consuming feed that contained CT had
increased DMI of a feed that contained terpenes (Mote et
al., 2007), and 4) in the current trial, J. ashei contained
greater CT than the CSH but had similar DMI (Table 3).
Terpenes can elicit olfactory cues (Müller-Schwarze,
1991; Straka, 1993; Bedoya-Pérez et al., 2014) and re-
duce herbivory of Juniperus trees (Markó et al., 2008;
Utsumi et al., 2009; Estell et al., 2014). Even though
Juniperus spp. contain volatile terpene oil, mechanical
drying, as done in this trial, can reduce total terpene con-
centration (Adams, 2010); a similar ground Juniperus
product contained less than 1% terpene oil (Stewart et
al., 2015). Others have reported that lamb DMI (similar
lambs and DDGS-based diets) was not affected when
mixed diets contained up to 36% ground J. pinchotii
4157
leaves and stems, which contained 0.43% terpene oil
(Whitney et al., 2014). Furthermore, DMI was similar
(P > 0.10) among all lambs during the preconditioning
period (data not shown) and DMI of lambs fed MESQ
was reduced during the first 14 d, even though terpene
concentrations are practically devoid in mesquite wood
(Papiez et al., 2009; Henciya et al., 2017). These results
and those of previous feeding trials, along with absence
of neophobia during the transition period, suggested that
terpenes were not a contributing factor in reducing DMI.
No treatment × day interaction (P = 0.28) was ob-
served for lamb ADG. When averaged across all days,
lambs fed CSH tended to have greater (P < 0.10) ADG
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than lambs fed ERC and had greater (P < 0.05) ADG
than lambs fed MESQ. Average daily gain differences
were likely due to less leaf material of ERC trees vs.
the other juniper species and due to the mature mes-
quite trees being harvested without leaves. Juniper
leaves are as digestible as alfalfa (Whitney and Muir,
2010) and, therefore, enhanced DE intake.
A day effect (P < 0.001) was observed for lamb
ADG: d 14 vs. 28 (P = 0.84), d 28 vs. 43 (P = 0.79), and
d 43 vs. 57 (P < 0.001). Due to the percentage of struc-
tural fiber decreasing in Period 2 diets (30% CSH or
ground wood in Period 1 and 14% CSH or ground wood
in Period 2), lamb ADG was greater at the end of Period
2 than at the end of Period 1 (d 28 to 57; P < 0.001),
which further supports the preceding DMI discussion
in regards to low-quality roughages causing dietary en-
ergy dilution. Lambs fed CSH had similar (treatment ×
day, P = 0.37; treatment effect, P = 0.66) G:F through-
out the trial compared with lambs fed wood-based diets.
A day effect (P < 0.001) was observed for lamb G:F: d
14 vs. 28 (P = 0.08), d 28 vs. 43 (P = 0.64), and d 43
vs. 57 (P < 0.001). Lamb G:F was greater at the end
of Period 2 than at the end of Period 1 (d 28 to 57; P <
0.001), likely due to greater dietary energy density.
No treatment × day interaction (P = 0.71) or treat-
ment effect (P = 0.12) was observed for lamb BW
throughout the trial (Fig. 2); however, a day effect (P <
0.001) was detected. At the end of the trial, lambs fed
RED, BLUE, ONE, ERC, and MESQ weighed 3.3, 2.6,
2.1, 6.3, and 3.9 kg less, respectively, than lambs fed
CSH. If ADG continues to be similar to the final 14 d on
feed, lambs fed RED, BLUE, ONE, ERC, and MESQ
would require an additional 7, 6, 4, 13, and 9 d, respec-
tively, in the feedlot to reach the final BW of lambs fed
CSH. The economic benefit of the extra days on feed
will be determined mainly by feed costs, which affects
cost per kilogram of feed. During Period 1, cost of feed
per kilogram of BW gain for lambs fed CSH, RED,
BLUE, ONE, ERC, or MESQ was $0.95, $1.00, $0.93,
$1.04, $0.97, and $1.02, respectively. During Period
2, cost of feed per kilogram of BW gain for lambs fed
4158 Whitney et al.

Table 4. Effects of replacing cottonseed hulls (CSH) with ground woody products on lamb rumen fluid profile1
Diet2 P-value4
Item CSH RED BLUE ONE ERC MESQ SEM3 T D T×D
pH 0.71 <0.001 0.15
d 14 6.30 6.35 6.37 6.29 6.50 6.42 0.10
d 57 6.12 6.13 6.18 6.04 6.08 5.90 0.09
Ammonia N, mg/dL 0.02 0.003 0.04
d 14 2.14 4.91 3.54 4.74 4.91 6.75* 0.71
d 57 6.10 7.43 6.44 3.89 5.83 6.48 0.99
Total VFA, mmol 0.03 <0.001 0.99
d 14 16.2 33.5 31.1 25.8 27.8 34.0 5.5
d 57 34.5 50.5 44.5 43.0 47.9 50.8 5.0
d 14 and 57 25.3 42.0* 37.8 34.4 37.8 42.4* 4.0
Acetate:propionate ratio 0.93 <0.001 0.04

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d 14 1.7 1.5 1.7 1.6 1.9 2.0 0.2
d 57 1.1 1.2 1.3 1.1 1.0 0.9 0.1
Acetate, mol/100 mol 0.007 <0.001 0.91
d 14 45.3 36.8 38.3 37.8 38.9 35.3 2.1
d 57 32.0 28.7 30.6 29.1 30.1 27.2 1.7
d 14 and 57 38.1 32.5* 34.2 33.1 34.2 31.0* 1.3
Propionate, mol/100 mol 0.22 0.005 0.01
d 14 28.1 26.5 23.5 24.2 20.7 18.7* 2.4
d 57 29.1 25.5 25.1 27.7 30.3 29.7 2.1
Butyrate, mol/100 mol 0.07 <0.001 0.01
d 14 18.1 20.0 27.2* 27.4* 22.8 20.9 2.1
d 57 13.2 1.36 13.6 10.9 11.7 11.1 1.2
1During Period 1 (d 0 to 27), lambs were fed a 70% concentrate diet and blood serum collected on d 27. Lambs were transitioned over 4 d into Period 2
(d 28 to 57) onto an 86% concentrate diet and blood serum collected on d 57.
2Treatment diets were nonagglomerated and ingredient composition differed only by roughage source; either CSH or ground woody products (redberry
juniper [RED; Juniperus pinchotii], blueberry juniper [BLUE; Juniperus ashei], one-seeded juniper [ONE; Juniperus monosperma], eastern red cedar
[ERC; Juniperus virginiana], or honey mesquite [MESQ; Prosopis glandulosa]). Juniperus (entire aboveground biomass) and Prosopis (entire aboveg-
round biomass except for leaves) species were chipped, dried, and hammer milled to pass a 4.76-mm sieve.
3SEM represents the greatest SEM.
4T = treatment effect; D = day effect.

Within row: *P < 0.05.

CSH, RED, BLUE, ONE, ERC, or MESQ was $0.82,
$0.76, $0.85, $0.76, $0.84, and $1.02, respectively.
Rumen Fluid Profiles
Lamb rumen fluid profile data are presented in
Table 4. No treatment × day interaction (P = 0.15) or
treatment effect (P = 0.71) was observed for ruminal
pH. Considering that lamb ruminal pH can increase
when CSH replace other roughage ingredients (Hsu et
al., 1987; Zhang et al., 2010), the results suggested
that feeding the woody-based diets provided adequate
rumen buffering capacity. The dense particles in the
ground wood contain greater concentrations of lig-
nin in relation to traditional roughage ingredients.
As previously discussed for a similar woody product
(Whitney et al., 2014), lignin was likely acting as an
innate buffer (Van Soest et al., 1991), especially in the
ventral rumen, where the majority of material in the
woody-based diets would reside.
Rumen ammonia N was similar between lambs
fed CSH and lambs fed woody-based diets, except
on d 14 (treatment × day interaction, P = 0.04), when
lambs fed CSH had less (P < 0.05) ruminal ammonia
N than lambs fed MESQ. Rumen ammonia N concen-
tration can increase due to greater intake of CP, de-
gradable protein (Satter and Slyter, 1974; Huntington
and Archibeque, 2000), or both and can decrease due
to greater starch fermentation, a reduction in rumen
pH (Lana et al., 1998; Rémond et al., 2002), and CT
consumption (Yu et al., 1995). Condensed tannin in-
take was relatively similar between lambs fed CSH
and MESQ; however, considering that lambs fed
CSH consumed greater amounts of total CP, degrad-
able protein, greater rumen ammonia N is expected. In
contrast, 48-h IVDMD was greater for the CSH diet
than for the MESQ diet, and lambs fed the CSH diet
had greater DE intake, which would tend to reduce
ammonia N. Even though ammonia N concentrations
were less than what would be expected for lambs fed
 Ground woody products as roughage sources 4159

Table 5. Effects of replacing cottonseed hulls (CSH) with ground woody products on lamb blood serum profile1
Diet3 P-value5
Item2 CSH RED BLUE ONE ERC MESQ SEM4 T D T×D
Glucose, mg/dL 0.008 <0.001 0.008
d0 84.9 78.5 81.8 73.4* 79.3 78.4 2.1
d 14 87.3 82.0 81.4 76.7* 80.0† 76.6* 1.6
d 57 86.1 86.6 87.3 85.8 87.9 85.6 1.9
NEFA, mEq/L 0.83 0.05 0.26
d0 0.10 0.10 0.11 0.12 0.11 0.11 0.01
d 14 0.12 0.10 0.12 0.10 0.14 0.12 0.01
d 57 0.11 0.10 0.10 0.11 0.12 0.10 0.01
BHB, µmol/L 0.17 <0.001 0.01
d0 437.5 462.3 552.6 421.4 470.3 397.7 51.5
d 14 461.0 570.2 734.5† 631.4 657.8 592.9 70.3

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d 57 395.6 471.7 438.7 366.0 398.8 377.6 37.5
SUN, mg/dL 0.25 0.10 0.10
d0 16.1 18.3 20.2 21.6 21.9 21.7 0.98
d 14 18.8 20.9 22.4 23.2 21.4 21.1 2.09
d 57 20.3 18.9 17.7 21.4 21.0 18.6 1.84
Creatinine, mg/dL 0.62 0.03 0.22
d0 0.65 0.71 0.66 0.76 0.71 0.70 0.03
d 14 0.67 0.70 0.66 0.76 0.73 0.69 0.04
d 57 0.74 0.73 0.73 0.77 0.71 0.72 0.03
d 14 and 57 0.70 0.71 0.69 0.76 0.72 0.70 0.02
Bilirubin, mg/dL <0.001 <0.001 <0.001
d0 0.10 0.11 0.10 0.10 0.10 0.10 0.004
d 14 0.09 0.11 0.11 0.13 0.12 0.16 0.006
d 57 0.10 0.12 0.11 0.12 0.12 0.13 0.004
Albumin, g/dL 0.29 <0.001 0.28
d0 2.91 2.91 2.89 2.96 2.99 2.97 0.05
d 14 3.05 2.89 2.89 3.00 2.99 3.00 0.05
d 57 3.13 3.10 3.05 3.18 3.06 3.13 0.05
Globulin, g/dL 0.24 <0.001 0.27
d0 2.69 2.64 2.56 2.75 2.63 2.69 0.09
d 14 2.68 2.45 2.45 2.69 2.66 2.68 0.09
d 57 2.34 2.28 2.30 2.44 2.39 2.58 0.10
A:G ratio 0.73 <0.001 0.04
d0 1.09 1.10 1.13 1.08 1.14 1.11 0.04
d 14 1.14 1.19 1.18 1.12 1.12 1.13 0.03
d 57 1.34 1.37 1.33 1.31 1.28 1.22 0.05
TP, g/dL 0.10 0.06 0.43
d0 5.6 5.55 5.49 5.73 5.63 5.66 0.11
d 14 5.75 5.35 5.35 5.73 5.63 5.7 0.12
d 57 5.49 5.39 5.38 5.6 5.45 5.71 0.11
ALT, units/L 0.001 <0.001 0.20
d0 6.38 7.50 7.13 6.75 6.88 5.88 0.56
d 14 6.88 9.38 9.63 8.51 9.13 8.63 0.58
d 57 5.38 7.86 7.38 7.00 9.22 6.00 0.70
d 14 and 57 6.12 8.6* 8.5* 7.8 9.13* 7.3 0.50
AST, units/L 0.56 <0.001 0.18
d0 55.6 61.7 54.1 58.4 62.5 48.8 5.2
d 14 68.7 62.8 64.1 63.6 80.9 84.8 8.3
d 57 53.1 59.8 51.6 63.1 62.0 49.4 3.4
GGT, units/L 0.09 0.02 0.04
d0 53.4 50.5 49.3 54.9 54.0 47.3 2.8
d 14 57.4 46.3* 47.4† 53.6 54.3 46.9† 2.5
d 57 54.0 45.4 46.7 52.0 50.2 49.2 2.4

Continued
4160 Whitney et al.

Table 5. (cont.)
Diet3 P-value5
Item2 CSH RED BLUE ONE ERC MESQ SEM4 T D T×D
CPK, units/L 0.79 <0.001 0.69
d0 244.1 257.5 137.5 213.9 208.3 157.5 72.1
d 14 108.4 84.6 98.0 177.2 68.4 155.6 38.5
d 57 53.5 67.2 58.6 62.8 64.4 83.6 13.6
Ca, mg/dL 0.84 <0.001 0.88
d0 10.0 10.2 9.9 10.1 10.1 10.0 0.13
d 14 10.3 10.3 10.2 10.5 10.3 10.4 0.18
d 57 10.4 10.4 10.4 10.5 10.3 10.5 0.16
P, mg/dL 0.16 <0.001 0.03
d0 10.4 10.5 10.6 9.1 9.8 10.2 0.35
d 14 9.2 7.7 7.7 8.3 7.7 8.0 0.42

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d 57 9.6 9.3 9.0 8.2 8.6 8.9 0.42
Mg, mEq/L 0.32 <0.001 0.003
d0 2.19 2.20 2.17 2.10 2.24 2.14 0.05
d 14 2.32 2.29 2.21 2.25 2.19 2.08† 0.06
d 57 2.36 2.28 2.22 2.31 2.48 2.31 0.06
Cl, mEq/L 0.72 <0.001 0.008
d0 115.3 112.7 115.9 112.8 113.4 115.6 1.0
d 14 113.9 115.4 114.6 114.0 114.2 116.7 0.9
d 57 116.8 118.0 117.6 115.8 117.2 116.5 1.3
Na, mEq/L 0.95 <0.001 0.81
d0 152.8 150.9 153.4 148.9 150.1 150.7 0.8
d 14 151.4 151.3 152.0 150.9 149.9 151.4 1.2
d 57 153.7 153.6 154.1 151.2 152.7 152.7 1.0
d 14 and 57 152.5 152.5 153.0 151.1 151.3 152.1 0.64
K, mEq/L 0.12 0.87 0.78
d0 6.13 6.45 6.04 5.79 6.30 6.13 0.25
d 14 6.49 5.99 6.13 5.69 6.27 6.15 0.26
d 57 6.36 6.42 6.33 5.61 6.11 6.26 0.22
Na:K ratio 0.18 0.98 0.69
d0 25.2 23.9 25.6 25.8 24.1 24.9 1.0
d 14 23.5 25.5 25.1 26.8 24.1 25.1 1.0
d 57 24.3 24.1 24.6 27.1 25.2 24.7 0.9
1During Period 1 (d 0 to 27), lambs were fed a 70% concentrate diet and blood serum collected on d 27. Lambs were transitioned over 4 d into Period 2
(d 28 to 57) onto an 86% concentrate diet and blood serum collected on d 57.
2BHB = β-hydroxybutyrate; SUN = serum urea N; A:G = albumin:globulin; TP = total protein; ALT = alanine aminotransferase; AST = aspartate ami-
notransferase; GGT = gamma-glutamyl transferase; CPK = creatine phophokinase.
3Treatment diets were nonagglomerated and ingredient composition differed only by roughage source: either CSH or ground woody products (redberry
juniper [RED; Juniperus pinchotii], blueberry juniper [BLUE; Juniperus ashei], one-seeded juniper [ONE; Juniperus monosperma], eastern red cedar
[ERC; Juniperus virginiana], or honey mesquite [MESQ; Prosopis glandulosa]). Juniperus (entire aboveground biomass) and Prosopis (entire aboveg-
round biomass except for leaves) species were chipped, dried, and hammer milled to pass a 4.76-mm sieve.
4SEM represents the greatest SEM.
5T = treatment effect; D = day effect.

Within row: †0.05 > P < 0.10; *P < 0.05.

diets containing a minimum of 17% CP, efficiency of
microbial protein synthesis was not likely negative-
ly affected, except for lambs fed CSH during Period
1. Satter and Slyter (1974) suggested that maximum
growth rates of rumen bacteria occurred at around 5.0
mg ammonia N/dL but that limiting concentrations
probably occurred at closer to 2.0 mg/dL.
Averaged across both days (treatment × day interac-
tion, P = 0.99), lambs fed CSH had less (P < 0.05) total
VFA than lambs fed RED or MESQ. No treatment × day
interaction (P = 0.91) was observed for molar proportion
of acetate. However, acetate (mol/100 mol) was greater
(P < 0.05) in lambs fed CSH than in lambs fed RED
or MESQ when averaged across both days. Treatment ×
day interactions (P = 0.01) were observed for molar pro-
portions of propionate and butyrate. On d 14, the mo-
lar proportion of propionate was greater (P < 0.05) in
lambs fed CSH than in lambs fed MESQ, but no other
 Ground woody products as roughage sources
differences were observed. A treatment  × day interac-
tion (P = 0.04) was observed for the acetate:propionate
ratio, but no differences (P > 0.10) were observed within
day. On d 14, the molar proportion of butyrate was less
(P < 0.05) in lambs fed CSH than in lambs fed BLUE
or ONE, but no other differences were observed. The
reason for greater total VFA and less molar proportion
of acetate in lambs fed MESQ or RED, compared with
lambs fed CSH, is not easily discerned.
Blood Serum Profiles
Lamb blood serum profiles are presented in
Table 5. Treatment × day interactions (P < 0.05) were
observed for serum glucose, BHB, total bilirubin, and
the albumin:globulin ratio. Lambs had similar (treat-
ment × day and treatment effect, P > 0.10) serum
NEFA, serum urea N, creatinine, albumin, globulin,
and total protein. On d 14, lambs fed CSH had great-
er (P < 0.05) serum glucose than lambs fed ONE or
MESQ and tended to have (P < 0.10) greater serum
glucose than lambs fed ERC. Lambs fed BLUE tended
to have greater (P < 0.10) serum BHB than lambs fed
CSH on d 14; however, BHB concentrations less than
1,000 µmol/L do not indicate ketosis (Smith and Davis,
2015). Within day, lambs had similar (P > 0.10) serum
total bilirubin and albumin:globulin ratio.
A treatment × day interaction (P = 0.04) was ob-
served for serum gamma-glutamyl transferase (GGT).
On d 14, lambs fed CSH had greater (P < 0.05) serum
GGT than lambs fed RED and tended to have (P < 0.10)
greater GGT than lambs fed BLUE or MESQ. Serum
GGT functions as a clinical indicator of hepatic disor-
ders (Cornelius, 1989; Stämpfli and Oliver-Espinosa,
2015). No treatment × day interaction (P = 0.20) was
observed for serum ALT. However, when averaged
across d 14 and 57, lambs fed CSH had less (P < 0.05)
serum ALT than lambs fed RED, BLUE, or ERC. The
normal reference value for serum ALT ranges from 22
to 38 units/L (Stämpfli and Oliver-Espinosa, 2015),
which is greater than any of the values reported in the
present trial. Lambs had similar (treatment × day and
treatment effects, P > 0.10) concentrations of serum
aspartate aminotransferase and creatine phophokinase.
Treatment × day interactions (P < 0.10) were ob-
served for serum P, Mg, and Cl, but no differences
(P > 0.12) were detected within day for serum P or Cl.
Within d 14, serum Mg tended to be greater (P < 0.10)
in lambs fed CSH than in lambs fed MESQ; however,
no lambs experienced hyomagnesemia as described by
others (McFarlane and Fleming, 2015; Stämpfli and
Oliver-Espinosa, 2015). When collectively evaluated,
serum chemistry results did not indicate that using up
to 30% ground woody products in lamb feedlot diets
4161
negatively affected animal health and, more specifi-
cally, cause hepatotoxicity.
Conclusions
Results suggested that even though lamb DMI
was reduced during Period 1 when growing diets con-
tained 30% J. pinchotii, J. monosperma, J. virginiana,
and P. glandulosa, rumen fluid parameters were not
negatively affected and all lambs fed woody-based
diets remained healthy throughout the trial. Due to
the physical characteristics of the woody-based diets,
lamb growth performance would probably increase if
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a small amount of ground hay was included. Ground
woody products are unique in that they are not subject
to seasonal feed pricing and availability, require no in-
puts by man to establish and grow, and are the only
feed ingredients that can increase rangeland forage
production after harvest. Therefore, the development
of a commercialized industry to supply ground woody
products for livestock production is warranted.
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