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1982) PARFITT, et al: POTATO VINE SILAGE 415

T H E N U T R I T I O N A L VALUE OF PRESSED P O T A T O VINE S I L A G E 1

Dan E. Parfitt, S.J. Peloquin and N.A. Jorgensen 2

Abstract

Potato (Solanum tuberosum L.) vines were studied to determine if they


could be successfully ensiled and fed to ruminant animals. Russet Burbank
potato vine and alfalfa (Medicago sativa L.) silages were fed to goats
(Capra hircus) to measure in vivo digestibility and to determine if potato
glycoalkaloids were aversive or toxic. No differences in the feeding patterns
between control and experimental animals were observed. Alfalfa silage was
more digestible than the Russet Burbank potato vine silage due to a higher
content of digestible cell wall materials. Digestibility differences between
potato clones and harvest dates were found when in vivo digestibility tests
were performed. W729R was the most digestible clone studied, especially
for the early harvest dates. Most of the clones had vines that were more
digestible, in vitro, than alfalfa. Vine glycoalkaloid contents of the experi-
mental clones were similar to the values reported in the literature. There was
no apparent correlation between vine glycoalkaloid content and percent di-
gestible dry matter. A positive correlation of 0.38 between digestible dry
matter and dry vine yield was found. Potato vine silage, produced after
pressing the vines to remove water, was an acceptable feed for ruminant
animals. Additional in vivo testing is necessary to determine if toxic effects
due to glycoalkaloids or pesticides from long term feeding will be a problem.

Resumen

Se estudi6 el rastrojo de la papa (Solanum tuberosum L.) para determi-


nar si es-factible ensilarlo con 6xito y utilizarlo para alimentar rumiantes. A
grupos de cabras (Capra hircus) se les di6 ensilaje de alfalfa (Medicago sati-
va L.) y de rastrojo de papa Russet Burbank para medir digestibilidad en
vivo y determinar si los glicoalcaloides de la papa eran t6xicos o causaban
aversi6n. No se encontraron diferencias en los h~bitos de alimentaci6n entre
los testigos y los tratamientos. El ensilaje de alfalfa fue mhs digestible que el
de rastrojo de papa, debido a un contenido mayor de pared celular digestible.

~Research supported by the Collegeof Agricultural and Life Sciences, Universityof Wisconsin,
grants from the National Science Foundation (PCM-77-24330) and the International Potato
Center, and a gift from Frito-Lay, Inc.
ZFormer graduate student and Professor, Departments of Horticulture and Genetics, and Pro-
fessor, Department of Dairy Science, University of Wisconsin, Madison, WI 93706. Present
address of D.E.P.: PomologyDepartment, University of California, Davis, CA 95616.
Received for publication March 4, 1982.
KEY WORDS: Feeding value, haulm, potato, nutrition, ruminant animals, silage, Solanum
tuberosum L., vine.
416 AMERICAN POTATO JOURNAL (Vol. 59

Cuando se hicieron pruebas de digestibilidad en vivo, se encontraron dife-


rencias en digestibilidad entre clones de p a p a y edades de corte. E1 W729R
fue el mhs digestible entre los clones estudiados, especialmente cuando el
corte se hizo a edad temprana. La mayoria de los clones tenia rastrojo que
era m~ts digestible, in vitro, que la alfalfa. Los contenidos de glicoalcaloides
del rastrojo fueron similares a aqu611os referidos en la literatura. No hubo
correlaci6n clara entre los glicoalcaioides del rastrojo y el porcentaje de
materia seca digestible. Se encontr6 una correlaci6n positiva de 0.38 entre
materia seca digestible y producci6n de rastrojo seco. El ensilaje de rastrojo
de papa, producido despu6s de comprimir el rastrojo para extraerle el agua,
fue un alimento aceptable para rumiantes. Se reguieren pruebas adicionales
en vivo para determinar si se presentaran problemas debido al afecto de los
glicoalcaloides o los pesticidas en periodos largos de alimentaci6n con dicho
ensilaje.
Introduction
Potatoes (Solanum tuberosum L.) produce large quantities of vines
that are not presently utilized (7). One possible use for these vines is to ensile
them and feed the silage to livestock. Unfortunately, potato vines are usually
too high in moisture to make good silage. With the development of alfalfa
pressing machinery, the possibility now exists for removing excess water
from the vines and successfully ensiling them (8). The nutritional value of
forages is a function of three parameters: digestibility, voluntary intake,
and palatability (10). A forage should not contain toxic materials and should
have the necessary constituents for animal growth.
Several methods have been developed to measure the a m o u n t of digest-
ible material in forages. A c o m m o n in vivo measure of digestibility is the
total digestible nutrients (TDN) procedure. T D N is determined by compar-
ing the amounts of plant material in the feed and feces. Several in vitro pro-
cedures have been developed that are more convenient than in vivo methods.
In vitro procedures permit rapid measurement of a large n u m b e r of samples.
The Tilley-Terry in vitro digestibility procedure is frequently used. Results
from in vivo and in vitro procedures are highly correlated, with r = 0.88 to
0.98 (1).
Several chemical assays are available for determining the nutritional
value of forages. Three assays that are often used are the cell wall constitu-
ents (CWC), acid detergent fiber (ADF), and acid detergent lignin (ADL)
procedures. C W C is a measure of total fiber (does not contain soluble cel-
lular materials); ADF is a measure of lignin, cellulose, and ash fractions;
and A D L is a measure of lignin and ash (11). Estimates for cellulose and
hemicellulose content are derived from these assays by subtraction. Soluble
nutrients are digestible; the cell wall components are partially digested by
ruminants. Celluloses are digested more slowly than hemicelluloses or solu-
ble sugars. Lignin is indigestible, and may be related to in vivo digestibility (1).
1982) PARFITT, et al: POTATO VINE SILAGE 417

Voluntary intake is an important forage parameter that can affect the


weight gain of ruminant animals. Expressed as a percentage of digestible in-
take for alfalfa, voluntary intake was responsible for 47°7o and digestibility
accounted for 26°7o (11). Voluntary intake is positively correlated with di-
gestibility. The rate at which a forage is digested and passes through a rumi-
nant animal's digestive system may affect the voluntary intake of the forage.
Voluntary intake levels of a given forage can vary a m o n g animal species and
between animals within a species (3, 5). A positive correlation between palat-
ability and voluntary intake was found in a number of studies (5). Palatabil-
ity may not be an important concept if only one type of feed is available to
the animal. That would usually be the situation when animals are fed silage
in feedlots.

Materials and Methods


Four male castrated goats (Capra hircus) were used in a total collection
digestion trial with a 14 day adjustment period followed by a seven day col-
lection period to evaluate pressed and ensiled Russet Burbank potato vines,
pressed alfalfa (Medicago sativa L.) silage, and conventional alfalfa silage
control. Two of the goats were fed ensiled Russet Burbank potato vines (har-
vested 112 days after planting in 1976), for three weeks. The potato vines
were pressed with a screw press to reduce the moisture content of the vines
from 90°70 to 50°7o before ensiling. This experiment permitted evaluation of
digestibility, cell wall constituents, voluntary intake, and acute toxicity of
the vines.
L a b o r a t o r y tests for the nutritional value of potato vine cultivars at
three harvest dates in 1976 (105, 120, and 135 days from planting) were car-
ried out in 1977 and 1978. CWC, ADF, and A D L procedures were performed
to determine the amounts of soluble nutrients, hemicellulose, cellulose, and
lignin. Vines from the 1976 yield trial were assayed for total protein content,
in vitro digestible dry matter, and glycoalkaloid concentration. Two hundred
grams to 500 g vine samples f r o m each clone and harvest date were dried at
80°C for 48 hours, and were ground to pass through a 0.425 m m (40 mesh)
screen. Total protein content was determined by a semi-micro Kjeldahl pro-
cedure.
In vitro digestible dry matter assays were performed via the Tilley-Terry
in vitro rumen fermentation procedure. One forth gram samples of dried
vine were placed in 50 ml centrifuge tubes with 20 ml of a buffer solution
and five mililiters of rumen fluid from a cow. Stoppers were placed in the
tubes after the tubes were flushed with CO2 (the reaction is anaerobic). The
tubes were incubated for 48 hours at 39°C and the mixtures were filtered.
The solid residues were dried and weighed. The amounts of digestible dry
matter (DDM) were calculated by subtracting the residue weights from the
initial sample weights. Controls and dehydrated alfalfa standards were used
418 AMERICAN POTATO JOURNAL (Vol. 59

to adjust the results to a c o m m o n base. Four harvest dates (90, 105, 120,
and 135 days from planting) were analyzed.
Glycoalkaloid assays were performed on vine samples from the 135 day
harvest o f 1976. The extraction procedure was a modification o f a bisolvent
extraction method (12). One half gram vine samples were extracted with 15
ml o f hot 2:1 methanol-chloroform solution for three hours. Ten milliliters
of 0.006 N HC1 was added to each tube. The samples were heated to 70°C to
flocculate the glycoalkaloids. The flocculated materials were washed twice
with 25 ml of dilute ammonium hydroxide and dried. The precipitates were
dissolved in 15 ml of 100% methanol. Ten milliliter aliquots of the methanol
solutions were assayed for total glycoalkaloid concentration. A nonaqueous
acid-base titration procedure (2) was modified slightly and used to determine
TGA concentrations in the aliquots.
Factorial variance analyses were used to analyze the protein and digest-
ibility data from the 1976 potato vine samples. A randomized complete block
variance analysis was performed on the glycoalkaloid data. Covariance
analyses were performed to permit the calculation of treatment correlations
for dry weight vine yield vs. digestibility, and glycoalkaloid concentration
vs. digestibility.
Results

The results of the feeding and digestion study are presented in Tables 1
and 2. Results are presented for normally ensiled alfalfa and pressed alfalfa
silage, as well as pressed potato vine silage. The goats consumed quantities

TABLE 1. - - In vivo digestibility o f ensiled alfalfa and Russet Burbank


potato vines.

Weof dry wt.


Pressed potato vine Pressedalfalfa Regularalfalfa
Digestible dry matter 53.1 60.4 60.9
Digestible protein 60.3 64.3 69.6
Digestible cell wall 53.4 61.6 56.8

TABLE 2. - - Chemical analysis o f alfalfa and Russet Burbank potato


vine silages.

% of dry wt.
Pressed potato vine Pressed alfalfa Regularalfalfa
Crude protein 11.4 15.2 18.5
CWC 60.6 62.7 52.5
ADF 54.9 49.0 40.7
HemiceUulose 5.7 13.6 11.8
Cellulose 37.3 40.0 33.1
Lignin + ash (ADL) 10.6 9.0 7.6
1982) PARFITT, et a# POTATO VINE SILAGE 419

o f p o t a t o vine silage t h a t were c o m p a r a b l e tO the a l f a l f a silage. T h e g o a t s


e x h i b i t e d n o toxic r e a c t i o n s f r o m c o n s u m i n g the p o t a t o vines.
T h e p o t a t o vine silage was s o m e w h a t lower in digestible d r y m a t t e r (in
vivo), p r o t e i n , a n d cell walls t h a n the a l f a l f a silage ( T a b l e 1). T h e p o t a t o
vine silage c o n t a i n e d less p r o t e i n a n d hemicellulose t h a n the p r e s s e d a l f a l f a
silage, while the pressed a l f a l f a silage h a d less p r o t e i n a n d m o r e cell wall
c o m p o n e n t s t h a n the n o r m a l a l f a l f a silage (Table 2). T h e q u a n t i t i e s o f lignin
were s o m e w h a t higher in the pressed p o t a t o vines t h a n in t h e pressed
a l f a l f a . L i g n i n c o n t e n t o f n o r m a l l y ensiled a l f a l f a was lower t h a n in the
silages t h a t were pressed. L i m i t e d o b s e r v a t i o n n u m b e r s p r e c l u d e d statistical
a n a l y s i s o f the d a t a in T a b l e I a n d 2.
T h e r e were significant in v i t r o digestible d r y m a t t e r d i f f e r e n c e s for vines
b e t w e e n clones, dates, a n d specific clone × d a t e c o m b i n a t i o n s (Tables 3 a n d
4). I n v i t r o digestibility values were g e n e r a l l y v e r y high f o r the p o t a t o vines.

TABLE 3. - - F values f o r protein content, % digestible dry matter, and


vine glycoalkaloid content o f potato vines harvested in 1976.

Source Protein Digestible Glycoalkaloid


content dry matter content
Block 2.22 0.41 0.82
Clone 1.27 4.92"* 3.12"
Date 32.49** 11.98**
Clone x date 0.60 1.94*

*, ** -Significant at the 5% and 1% level, respectively.

TABLE 4. - - M e a n in vitro % digestible dry matter f o r potato vines


harvested in 1976.

Days after planting


Means for
Clone 90 105 120 135 clones
Kennebec 73.8 76.2 71.8 71.4 73.3
Russet Burbank 73.7 72.1 73.8 67.1 71.7
Superior 74.4 74.5 71.3 71.4 72.9
W729R 80.6 75.7 73.4 73.2 75.7
PSDT 5 77.1 73.0 71.6 73.1 73.7
PSDT 17 77.2 72.4 69.3 67.6 71.6
PSDT 24 72.3 73.6 70.2 74.8 72.7
PSDT 41 71.0 69.8 72.4 65.2 69.6
Means for dates 75.0 73.4 71.7 70.5
Alfalfa standard = 69.2

5% LSD for clones = 2.3


5% LSD for dates = 1.6
5% LSD for clonex date:4.6
420 AMERICAN POTATO JOURNAL (Vol. 59

Only three clone × date combinations had less digestible dry matter than the
alfalfa standard. The potato vines contained the most digestible dry matter
early in the season. W729R, P S D T 5, and Kennebec were the most digest-
ible clones, while PSDT 41, Russet Burbank, and P S D T 17 were the least
digestible clones.
Differences between the protein content of potato vines harvested on
different dates were observed (Table 3). The vine protein content of all
clones declined later in the growing season. Mean vine protein contents of
17.0°70, 15.0070, and 11.4070 were found at 105,120, and 135 days from plant-
ing, respectively. The 5070 LSD for these values was 1.4%.
Pressed potato vines were tested for protein content to determine whether
pressing the juice from the vines lowered the protein content o f the ensilable
forage. Chopped vines (112 day harvest) had a protein content o f 18.0070 _+
0.1%0 and pressed vine material had a protein content of 11.0070 + 0 . 1 % .
These results are in agreement with the analysis from the feeding study.
Protein values for Russet Burbank pressed vines were almost identical
(11.0070 vs. 11.4%) for both analyses. Protein levels for alfalfa silage de-
creased f r o m 18.50/0 to 15.2% with pressing. Thus, significant amounts of
protein were lost from the forage during the water removal procedure.
The F values and means for vine glycoalkaloid content (TGA) from
clones at 135 days from planting are presented in Tables 3 and 5. The last
harvest date was selected for testing because that was the harvest date that
TABLE 5. - - Mean total glycoatkaloM (TGA) concentrations for potato
vines
harvested 135 days from planting in 1976.
Clone TGA
mg/100 g dry vine
Kennebec 105
Russet Burbank 246
Superior 163
W729R 207
PSDT 5 88
P S D T 17 100
P S D T 24 286
P S D T 41 295

5% LSD = 145

was considered to be the most likely date for economical potato vine har-
vesting. Considerable variation in T G A content was present a m o n g clones.
PSDT 5, P S D T 17, and Kennebec had the lowest T G A concentrations.
PSDT 41, P S D T 24, and Russet Burbank had the highest T G A concentra-
tions.
A significant positive correlation was observed between in vitro digest-
ible dry matter and vine yields (dry weight). The unbiased correlation coef-
1982) PARFITT, et al: POTATO VINE SILAGE 421

ficient was 0.38_+ (0.20 to 0.53) for P = 0 . 0 5 . The correlation between in


vitro digestibility and T G A content was nonsignificant. The unbiased corre-
lation coefficient was - 0 . 3 8 + ( - 0 . 6 7 to 0.00) for P = 0.05.
Discussion
The feeding study demonstrated that potato vine silage was an accept-
able feed for ruminant animals. The study was not a test of palatability,
since the animals were not offered a choice of feeds. The test indicated that
voluntary intake levels for potato vine silage were similar to voluntary intake
levels for alfalfa silage. Potato vines are quite digestible. Although the in
vivo and in vitro assays gave different results for potato vine vs. alfalfa di-
gestibility, the high digestibility of potato vines was adequately demonstrated.
Potato vine digestibility declined late in the growing season, but was high
enough to provide good nutritive value at the later harvest dates. Potato
glycoalkaloids did not appear to be detrimental to the animals. Similar re-
suits were reported by Nicholson, et al. (6) for sheep. The glycoalkaloids
were apparently degraded during the ensiling process or in the animals'
rumens. Acidic systems are known to cleave the glycosides from the
aglycones. Without attached sugars, the alkaloids are much less soluble and
may be less biologically active. Rumen bacteria may also be capable of de-
grading the aglycones. Nicholson's study provided evidence that ensiling
also reduced the amount of glycoalkaloid in the vines.-Glycoalkaloid con-
centrations were reduced by 30% to 50°-/o in that study.
A significant amount of glycoalkaloid was present in the potato vines.
Nicholson, et al. (6) found T G A levels in Kennebec and Russet Burbank
vines (184 mg and 224 mg/100 g o f dry vine) that were similar to the values
reported in the present study. Street, et al. (9) found T G A levels of 13 m g /
100 g of fresh leaves (equal to 130 mg/100 g of dry leaves) for King Edward
potatoes in late August. Wolf and Duggar (13) observed T G A levels of
about 200 mg/100 g of dry leaves and stems for Russet Burbank. The results
from these potato vine glycoalkaloid studies should be regarded as approxi-
mations of the true T G A levels in potato vines. These T G A values and values
presented in Table 5 may have systematic errors due to incomplete recovery
of the glycoalkaloids. Therefore, T G A comparisons between specific clones
should be made with caution. T G A levels declined during the growing season
in all of the cited studies. Harvesting potato vines later in the season could
minimize the amount of glycoalkaloid in the vines. Economic and yield con-
siderations also favor late harvest dates. Glycoalkaloids do not appear to be
a problem when potato vines are ensiled and used as feed for ruminant
animals.
Significant toxic, carcinogenic, or tetratagenic effects, from fungicides
or pesticides that are applied to the crop, could be a problem when potato
vines are fed. Research pertaining to the effects of agricultural chemicals on
ruminant animals has not been reported for potatoes.
422 AMERICAN POTATO JOURNAL (Vol. 59

P r o t e i n c o n c e n t r a t i o n in p o t a t o vines declined to m a r g i n a l l y a c c e p t a b l e
levels b y 135 days f r o m p l a n t i n g . P r o t e i n c o n c e n t r a t i o n g r e a t e r t h a n ten
p e r c e n t is c o n s i d e r e d to be a d e q u a t e for r u m i n a n t f o r a g e s (4). Vine p r o t e i n
c o n t e n t was r e d u c e d b y pressing a l f a l f a o r p o t a t o vines. T h e r e f o r e , a
n i t r o g e n s u p p l e m e n t m a y be r e q u i r e d with pressed p o t a t o vines to s u p p o r t
high levels o f a n i m a l p e r f o r m a n c e .
P o t a t o vines s h o u l d be a g o o d f o r a g e , especially i f s u p p l e m e n t a l n i t r o -
gen is a d d e d t o the silage. A l f a l f a m a y be m o r e n u t r i t i o u s t h a n p o t a t o vines,
b u t d o e s n o t p r o v i d e a high v a l u e v e g e t a b l e c r o p ( p o t a t o e s ) at the s a m e
time. B r e e d i n g for n u t r i t i o n a l q u a l i t y c o u l d be l i m i t e d to testing the best
clones f r o m yield trials for in v i t r o digestibility. C l o n e s with digestible vines
will p r o b a b l y be f o u n d if this p r o c e d u r e is f o l l o w e d , since vine yield (dry
wt.) a n d digestibility are positively c o r r e l a t e d . Breeding f o r p r o t e i n q u a n t i t y
is n o t r e c o m m e n d e d , as no genetic v a r i a b i l i t y was o b s e r v e d . G l y c o a l k a l o i d
screening is n o t feasible due to the l i m i t e d reliability o f the r a p i d a s s a y p r o -
cedures t h a t a r e presently a v a i l a b l e a n d the q u e s t i o n a b l e significance o f
g l y c o a l k a l o i d s as q u a l i t y f a c t o r s in silage. T h e final test for a n y p o t a t o clone
t h a t is selected for release to g r o w e r s s h o u l d be a large scale feeding trial to
e s t i m a t e the f o r a g e p a r a m e t e r s o f digestibility, v o l u n t a r y i n t a k e , p a l a t a b i l -
ity, a n d l o n g t e r m toxic effects u n d e r c o m m e r c i a l c o n d i t i o n s .

Acknowledgments
I wish to t h a n k Dr. R i c h a r d K o g e l f o r pressing the p o t a t o vines a n d
P a u l F r i t s c h e l f o r helping set u p the digestibility assays.

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2. Fitzpatrick, T.J., and S.F. Osman. 1974. A comprehensive method for the determination of
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10. Waldo, D.R. 1970. Factors influencing voluntary intake of forages. Proc. Nat. Conf. For-
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