![In situ dry matter disappearance (DMD) and in vitro dry matter digestion (IVDMD) of kudzu (P. montana) plant parts [20].](https://www.researchgate.net/publication/336454164/figure/tbl2/AS:813021198811136@1570850767022/In-situ-dry-matter-disappearance-DMD-and-in-vitro-dry-matter-digestion-IVDMD-of-kudzu_Q320.jpg)
![Summary of qualitative tests from phytochemical screening and analysis of kudzu (Pueraria phaseoloides) [43].](https://www.researchgate.net/publication/336454164/figure/tbl3/AS:813021198819328@1570850767049/Summary-of-qualitative-tests-from-phytochemical-screening-and-analysis-of-kudzu-Pueraria_Q320.jpg)
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agriculture
Communication
A Review of Kudzu’s Use and Characteristics as
Potential Feedstock
Joseph P. Gulizia * and Kevin M. Downs
School of Agriculture, Middle Tennessee State University, Murfreesboro, TN 37132, USA;
kevin.downs@mtsu.edu
*Correspondence: jpg3y@mtmail.mtsu.edu; Tel.: +1-(615)-898-5217
Received: 23 August 2019; Accepted: 8 October 2019; Published: 11 October 2019
Abstract:
This review assesses the potential use of kudzu (Pueraria montana var. lobata) as a feedstock
for livestock. Kudzu in the United States is a recognized invasive plant species that has continued to
cause problems for the environment and land owners. In kudzu’s native countries, it has continued to
have beneficial uses beyond being an adequate form of soil erosion control. Never the less, kudzu is a
rampant weed that causes harm to many environments. In the United States, local farm owners have
used ruminant species as a form of biological control to prevent the spread of kudzu and provide
their animals with a high nutrition feed supplement. However, there are few reports that assess
ruminal degradability in ruminants and kudzu quality. There is great potential for kudzu as a feed
supplement for livestock species. Furthermore, using kudzu as a feed supplement for livestock
species serves a dual-purpose of biologically controlling the spread of kudzu while providing those
animals with a high-quality feedstuff.
Keywords: kudzu; feedstock; livestock; biological control; invasive species
1. Introduction
During its active growing season, kudzu (Pueraria montana var. lobata) rapidly engulfs many
woody and herbaceous areas in the Southeastern United States. It is an invasive plant species that has
persistently evaded a definitive control method, and, as such, has created land management problems
for many land owners. A major problem that land owners face is the destruction occurring from kudzu
on their desired forages for agricultural production. Biological control using ruminants, specifically
browsers, however, can be a beneficial control method for both the owner of the land and the animals.
Kudzu is known to have a high nutritive content that can benefit animals. There is, however, limited
data on kudzu’s ruminal degradability in ruminants. This review will focus on kudzu’s characteristics,
uses, and qualities and assess its use as a potential feedstock for livestock species.
2. History of Kudzu
There are 17 species of kudzu in the genus Pueraria throughout the world (Table 1), all of which
are native to China, Taiwan, Japan, and India [1–3].
Agriculture 2019,9, 220; doi:10.3390/agriculture9100220 www.mdpi.com/journal/agriculture
Agriculture 2019,9, 220 2 of 15
Table 1. Species of Pueraria throughout the world [3].
P. tuberosa P. lacei
P. sikkimensis P. bella
P. candollei P. pulcherrima
P. mirifica P. phaseoloides
P. lobata P. peduncularis
P. imbricata P. stricta
P. edulis P. wallichii
P. alopecuroides P. rigens
P. calycina
These cultures have implemented various uses of kudzu, including medicinal purposes or making
it into cloth and paper [
2
]. Kudzu was first introduced into the United States from Japan in 1876
as a display at the first official World’s Fair in Philadelphia, Pennsylvania in the Plant Exhibition
section [
2
,
4
]. During the late 19th century, kudzu’s broad leaves and dense growth was initially used as
shade for porches and courtyards in the Southern United States. Kudzu quickly became popular and
more common among Southern U.S. farmers for its advertised multi-purpose uses, including as soil
erosion control, a cheap forage for livestock, and various practical uses around their homes [
2
]. In the
1930s, the U.S. Natural Resource Conservation Service (NRCS) oversaw combating soil erosion from
improper agricultural practices [
2
]. They dispersed 85 million kudzu seedlings to Southern U.S. farmers
to establish kudzu plots for soil erosion control and land revitalization [
2
]. Kudzu use in soil erosion
control was optimal due to its rapid growth of vines that drop roots every few feet which grips and
prevents excess soil movement [
5
]. Land revitalization came through kudzu’s property of being a soil
nitrogen fixer, which refilled overused, nitrogen deficit soils. In the 1930s, the U.S. government offered
an $8 per acre incentive to plant kudzu seedlings. By 1946, there were approximately 1.21 million
hectares of kudzu in the Southeastern United States [2].
Kudzu began gaining negative attention by the early 1950s, as it spread rampantly throughout the
Southern U.S., causing problems for farmland owners [
2
]. It was destructive in killing trees, collapsing
buildings, and destroying utility poles by aggressively traveling up these structures and forming a
dense mass that would strain their integrity [
6
]. The climate of the Southeastern United States was
ideal for kudzu to thrive, leading to it being placed on the United States Department of Agriculture
(USDA) common weed list in 1970 [
2
]. Kudzu became an invasive plant species in the United States
because it had no natural competitors in the environment to regulate its growth, as it would in its
native Asian countries. In 1997, the U.S. Congress voted to place kudzu on the Federal Noxious Weed
List. There is now an estimated 2.83 million hectares of land in the Southeastern United States that has
been engulfed with this invasive plant species [2].
3. Characteristics of Kudzu
Kudzu (P. montana), the species that is predominately found in the Southern U.S., is a large,
trifoliate-leaved, semi-woody, perennial vine that belongs to the legume family [
2
,
5
,
6
]. Plant species of
the legume family are known for being soil nitrogen fixers. Kudzu vines can grow up to 0.3 m per
day in early summer and as much as 18 m total during the growing season (May–October) [
2
,
5
,
7
].
It spreads from the root crown in any direction and will root at the vine nodes every few feet to
establish new growths [
2
,
5
]. The spread rate of kudzu can be accelerated by small vines of other plants
because kudzu can consistently twine around smaller vines more swiftly than large tree trunks [
5
,
8
].
Its tuberous roots (descended at the nodes) help maintain a heavy carbon reserve. Roots can reach a
depth of 4 m and weigh as much as 91–136 kg in older kudzu patches [
2
,
5
,
9
]. The taproot is enlarged
and beneficial in that it aids the plant in survival during drought periods [4,5].
Asexual regeneration is a frequent and common method by which kudzu multiplies. This occurs
every few feet where nodes (the areas on the vine where leaves and roots branch) will send down roots
establishing new root crowns [
5
]. There are few fruiting pods that develop viable seeds during the
Agriculture 2019,9, 220 3 of 15
optimal growing season, but its vegetative reproduction continually takes place as the nodes establish
roots [
6
]. During kudzu’s third growing season after germination, seed production will initiate by
producing a purple flower in late July to September in the U.S., if in full sun [
5
,
6
]. When seedpods are
produced there are only 1–2 viable seeds, and these seed pods are only found on climbing vines [
2
,
5
].
A prolonged exposure to high summer temperatures and increased soil temperatures will accelerate
seed germination by affecting seed coat permeability [5,10]. Attempts to eradicate kudzu by burning
may also promote seed germination, where potential new growth would emerge after the burning
attempt [5,10].
Kudzu is found in many places in the United States and can grow in a wide range of soil types,
including sandy soils, acid soils, lime soils, lowlands with high water tables, in over-heavy subsoil,
and in areas where winter soil temperatures do not drop below
−
32
◦
C [
2
,
11
]. Kudzu can be found
in open fields, road sides, and near forest edges, but its spread is at its peak in open fields [
5
]. The
widespread distribution of kudzu in the United States is shown in Figure 1.
Agriculture2019,9,xFORPEERREVIEW3of16
initiatebyproducingapurpleflowerinlateJulytoSeptemberintheU.S.,ifinfullsun[5,6].When
seedpodsareproducedthereareonly1–2viableseeds,andtheseseedpodsareonlyfoundon
climbingvines[2,5].Aprolongedexposuretohighsummertemperaturesandincreasedsoil
temperatureswillaccelerateseedgerminationbyaffectingseedcoatpermeability[5,10].Attemptsto
eradicatekudzubyburningmayalsopromoteseedgermination,wherepotentialnewgrowthwould
emergeaftertheburningattempt[5,10].
KudzuisfoundinmanyplacesintheUnitedStatesandcangrowinawiderangeofsoiltypes,
includingsandysoils,acidsoils,limesoils,lowlandswithhighwatertables,inover‐heavysubsoil,
andinareaswherewintersoiltemperaturesdonotdropbelow−32°C[2,11].Kudzucanbefoundin
openfields,roadsides,andnearforestedges,butitsspreadisatitspeakinopenfields[5].The
widespreaddistributionofkudzuintheUnitedStatesisshowninFigure1.
Figure1.Distributionmapofkudzu(P.montana)intheUnitedStates[12].nodata;species
reported.
ThismapshowsthatkudzuhasspreadfromtheSouthernU.S.andhasacquiredalevelof
hardinesstoendurecolderanddryerclimates.Kudzucanenduredroughtandhightemperatures,
butwillnotthriveinwetsoilsandyoungvegetativegrowthwilldieinlowtemperatures[6].When
reachingtemperaturesbetween30°Cand35°C,theefficiencyofphotosynthesiswillbeaffectedby
increasingheat[5].Kudzuwillgrowinmanydifferentsoils,buttheoptimalsoiltypeisadeep,loamy
soil[2,4–6].ThemostaggressiveplotsofkudzuareintheSoutheasternU.S.,withitsoptimalclimates
wherewintersaremild,summertemperaturesriseabove27°C,annualprecipitationexceeds102cm,
andsandyloamsoilsarewidespread[4,5].
Otherfactorsthatcanaffectgrowthofkudzuislightavailabilityandthepreviousexistingnative
plantlife.Askudzustartstoencountershade,growthwilldwindle,whereasindirectsunlightthe
growthratecanincrease3‐fold[5].Kudzucontainsahighleafsurfacearea,especiallywhenclimbing
trees,whichenhancesthephotosyntheticcompetitionforlight[5].Kudzuisconsideredheavilyshade
intolerantinhavingthehighestlightrequirementoutoffivenative(Rhusradicans,Clematisvirginiana,
Smilaxrotundifolia,Vitisvulpina,andParthenocissusquinque‐folia)andthreeexotic(Puerarialobata,
Lonicerajaponica,andHederahelix)vinespeciesintheSoutheastU.S.[5,13].
Kudzudiffersamongspeciesaroundtheworld.Americankudzu,comparedtotheJapanese
counterpart,isdistinctlydifferentinhowitoverwinters.Kudzuisconsideredasemi‐woody
perennialbecauseofitsoverwinteringability[5].Overwinteringisaprocesswherevinesdevelop
thickbark,accumulateannualringsofvasculartissue,andattainadesirablestemdiameter,usually
around2cm[5,14].Americankudzuwillproducetheseoverwinteringstemsonlyonthevigorous,
climbingplants,whereastheJapanesestrainwillproducetheoverwinteringstemsontheportions
thatliejustabovetheground[5,14].AnadditionaldifferencewiththeNorthAmericancultivarsof
Figure 1.
Distribution map of kudzu (P. montana) in the United States [
12
].
Agriculture 2019, 9, x FOR PEER REVIEW 3 of 16
initiate by producing a purple flower in late July to September in the U.S., if in full sun [5,6]. When
seedpods are produced there are only 1–2 viable seeds, and these seed pods are only found on
climbing vines [2,5]. A prolonged exposure to high summer temperatures and increased soil
temperatures will accelerate seed germination by affecting seed coat permeability [5,10]. Attempts to
eradicate kudzu by burning may also promote seed germination, where potential new growth would
emerge after the burning attempt [5,10].
Kudzu is found in many places in the United States and can grow in a wide range of soil types,
including sandy soils, acid soils, lime soils, lowlands with high water tables, in over-heavy subsoil,
and in areas where winter soil temperatures do not drop below −32 °C [2,11]. Kudzu can be found in
open fields, road sides, and near forest edges, but its spread is at its peak in open fields [5]. The
widespread distribution of kudzu in the United States is shown in Figure 1.
Figure 1. Distribution map of kudzu (P. montana) in the United States [12]. no data; species
reported.
This map shows that kudzu has spread from the Southern U.S. and has acquired a level of
hardiness to endure colder and dryer climates. Kudzu can endure drought and high temperatures,
but will not thrive in wet soils and young vegetative growth will die in low temperatures [6]. When
reaching temperatures between 30 °C and 35 °C, the efficiency of photosynthesis will be affected by
increasing heat [5]. Kudzu will grow in many different soils, but the optimal soil type is a deep, loamy
soil [2,4–6]. The most aggressive plots of kudzu are in the Southeastern U.S., with its optimal climates
where winters are mild, summer temperatures rise above 27 °C, annual precipitation exceeds 102 cm,
and sandy loam soils are widespread [4,5].
Other factors that can affect growth of kudzu is light availability and the previous existing native
plant life. As kudzu starts to encounter shade, growth will dwindle, whereas in direct sunlight the
growth rate can increase 3-fold [5]. Kudzu contains a high leaf surface area, especially when climbing
trees, which enhances the photosynthetic competition for light [5]. Kudzu is considered heavily shade
intolerant in having the highest light requirement out of five native (Rhus radicans, Clematis virginiana,
Smilax rotundifolia, Vitis vulpina, and Parthenocissus quinque-folia) and three exotic (Pueraria lobata,
Lonicera japonica, and Hedera helix) vine species in the Southeast U.S. [5,13].
Kudzu differs among species around the world. American kudzu, compared to the Japanese
counterpart, is distinctly different in how it overwinters. Kudzu is considered a semi-woody
perennial because of its overwintering ability [5]. Overwintering is a process where vines develop
thick bark, accumulate annual rings of vascular tissue, and attain a desirable stem diameter, usually
around 2 cm [5,14]. American kudzu will produce these overwintering stems only on the vigorous,
climbing plants, whereas the Japanese strain will produce the overwintering stems on the portions
that lie just above the ground [5,14]. An additional difference with the North American cultivars of
no data;
Agriculture 2019, 9, x FOR PEER REVIEW 3 of 16
initiate by producing a purple flower in late July to September in the U.S., if in full sun [5,6]. When
seedpods are produced there are only 1–2 viable seeds, and these seed pods are only found on
climbing vines [2,5]. A prolonged exposure to high summer temperatures and increased soil
temperatures will accelerate seed germination by affecting seed coat permeability [5,10]. Attempts to
eradicate kudzu by burning may also promote seed germination, where potential new growth would
emerge after the burning attempt [5,10].
Kudzu is found in many places in the United States and can grow in a wide range of soil types,
including sandy soils, acid soils, lime soils, lowlands with high water tables, in over-heavy subsoil,
and in areas where winter soil temperatures do not drop below −32 °C [2,11]. Kudzu can be found in
open fields, road sides, and near forest edges, but its spread is at its peak in open fields [5]. The
widespread distribution of kudzu in the United States is shown in Figure 1.
Figure 1. Distribution map of kudzu (P. montana) in the United States [12]. no data;
This map shows that kudzu has spread from the Southern U.S. and has acquired a level of
hardiness to endure colder and dryer climates. Kudzu can endure drought and high temperatures,
but will not thrive in wet soils and young vegetative growth will die in low temperatures [6]. When
reaching temperatures between 30 °C and 35 °C, the efficiency of photosynthesis will be affected by
increasing heat [5]. Kudzu will grow in many different soils, but the optimal soil type is a deep, loamy
soil [2,4–6]. The most aggressive plots of kudzu are in the Southeastern U.S., with its optimal climates
where winters are mild, summer temperatures rise above 27 °C, annual precipitation exceeds 102 cm,
and sandy loam soils are widespread [4,5].
Other factors that can affect growth of kudzu is light availability and the previous existing native
plant life. As kudzu starts to encounter shade, growth will dwindle, whereas in direct sunlight the
growth rate can increase 3-fold [5]. Kudzu contains a high leaf surface area, especially when climbing
trees, which enhances the photosynthetic competition for light [5]. Kudzu is considered heavily shade
intolerant in having the highest light requirement out of five native (Rhus radicans, Clematis virginiana,
Smilax rotundifolia, Vitis vulpina, and Parthenocissus quinque-folia) and three exotic (Pueraria lobata,
Lonicera japonica, and Hedera helix) vine species in the Southeast U.S. [5,13].
Kudzu differs among species around the world. American kudzu, compared to the Japanese
counterpart, is distinctly different in how it overwinters. Kudzu is considered a semi-woody
perennial because of its overwintering ability [5]. Overwintering is a process where vines develop
thick bark, accumulate annual rings of vascular tissue, and attain a desirable stem diameter, usually
around 2 cm [5,14]. American kudzu will produce these overwintering stems only on the vigorous,
climbing plants, whereas the Japanese strain will produce the overwintering stems on the portions
that lie just above the ground [5,14]. An additional difference with the North American cultivars of
species reported.
This map shows that kudzu has spread from the Southern U.S. and has acquired a level of
hardiness to endure colder and dryer climates. Kudzu can endure drought and high temperatures,
but will not thrive in wet soils and young vegetative growth will die in low temperatures [
6
]. When
reaching temperatures between 30
◦
C and 35
◦
C, the efficiency of photosynthesis will be affected by
increasing heat [
5
]. Kudzu will grow in many different soils, but the optimal soil type is a deep, loamy
soil [
2
,
4
–
6
]. The most aggressive plots of kudzu are in the Southeastern U.S., with its optimal climates
where winters are mild, summer temperatures rise above 27
◦
C, annual precipitation exceeds 102 cm,
and sandy loam soils are widespread [4,5].
Other factors that can affect growth of kudzu is light availability and the previous existing native
plant life. As kudzu starts to encounter shade, growth will dwindle, whereas in direct sunlight the
growth rate can increase 3-fold [
5
]. Kudzu contains a high leaf surface area, especially when climbing
trees, which enhances the photosynthetic competition for light [
5
]. Kudzu is considered heavily shade
intolerant in having the highest light requirement out of five native (Rhus radicans,Clematis virginiana,
Smilax rotundifolia,Vitis vulpina, and Parthenocissus quinque-folia) and three exotic (Pueraria lobata,
Lonicera japonica, and Hedera helix) vine species in the Southeast U.S. [5,13].
Kudzu differs among species around the world. American kudzu, compared to the Japanese
counterpart, is distinctly different in how it overwinters. Kudzu is considered a semi-woody perennial
because of its overwintering ability [
5
]. Overwintering is a process where vines develop thick bark,
Agriculture 2019,9, 220 4 of 15
accumulate annual rings of vascular tissue, and attain a desirable stem diameter, usually around
2 cm [5,14]. American kudzu will produce these overwintering stems only on the vigorous, climbing
plants, whereas the Japanese strain will produce the overwintering stems on the portions that lie just
above the ground [
5
,
14
]. An additional difference with the North American cultivars of kudzu is that
they have limited seed production and are less likely to thrive outside the Southeast U.S. [5].
4. Uses of Kudzu
In China and Japan, kudzu roots are dried and used for medicinal purposes to cure an array of
common ailments [
15
]. Japan, during the 1700s, also attempted to utilize fiber from stems to make
grass-like cloth and paper, and also grinding kudzu into flour for use in baking [
2
,
15
]. Asian grocery
and health food stores still import kudzu flour to sell in the U.S. [
9
]. Other traditional uses of kudzu are
as fiber to stuffcushions and chairs, as a mosquito repellent when burned, and to produce a palatable
honey [
15
]. During the initial years that kudzu was introduced in the U.S., it was used as an ornamental
vine (which was appreciated for its grape-like fragrance) to shade many southern U.S. homes [2].
As previously discussed, kudzu (P. montana) was first introduced to the United States as a means
for erosion control, but was eventually considered a rampantly unstoppable vegetation that would start
to take over the Southeastern United States. Kudzu continues to be an efficient method of soil erosion
control on steep embankments, but there are more noninvasive species (e.g., tall fescue and bahiagrass)
used now to address this issue [
7
,
15
]. Being a legume, kudzu has a dual-purpose of hosting nitrogen
fixing bacteria that enrich the soil and is also a good source of nutrients when fed to herbivorous
livestock [15].
5. Kudzu Nutrient Composition and Degradability
Kudzu (P. montana) is often compared to alfalfa. Kudzu leaves have a high nutritive value
comparable to that of alfalfa (Medicago sativa), a common flowering plant used for grazing, hay, and
silage for ruminants and other domestic herbivores [16–18] (Table 2).
Table 2.
Nutrient composition of alfalfa hay and fresh kudzu (P. montana) leaves (dry matter (DM)) basis)
Alfalfa, Hay, Sun-Cured, Midbloom 1Kudzu Leaves, Fresh 2
Crude protein (CP), % 18.7 17.5
Neutral detergent fiber (NDF), % 46.0 48.1
Acid detergent fiber (ADF), % 36.9 38.2
Ca, % 1.37 0.7
K, % 1.56 1.0
Mg, % 0.35 0.3
Fe, mg/kg 224.60 162.3
1NRC [19], 2Corley et al. [20].
As noted in Table 3, kudzu (P. montana) (kudzu aerial part (fresh), kudzu leaves (fresh), and
kudzu hay) is a high quality legume feedstuff. Kudzu fed as an aerial part (fresh), leaves (fresh), or
hay can satisfy most nutrient requirements for various ruminant species (Table 4). Kudzu silage has
a high nutrient composition that would satisfy nutrient requirements for many ruminants (20.15%
dry matter (DM), 92.01% organic matter (OM), 20.09% crude protein (CP), 8.14% Ash, 57.10% neutral
detergent fiber (NDF), 38.32% acid detergent fiber (ADF), 8.25% Lignin) [
21
]. Kudzu leaves are higher
quality than alternative kudzu feed sources, and satisfies nutrient requirements for most domestic
ruminants [
22
]. Additionally, based on National Research Council’s TDN criteria, kudzu leaves are
considered a high quality legume forage [
22
]. Nutrient composition data indicates that kudzu has
substantial potential as a feedstock for ruminant livestock species.
Agriculture 2019,9, 220 5 of 15
Table 3.
Chemical and nutrient composition of kudzu (P. montana), aerial part, fresh, kudzu (P. montana)
leaves, fresh, and kudzu (P. montana) hay (DM basis) [20,23–31].
Item Kudzu, Aerial Part, Fresh Kudzu Leaves, Fresh Kudzu Hay
Avg ASD BMin CMax DNb EAvg SD Min Max Nb Avg
Dry matter, % as fed 26.5 5.5 17.4 35.0 8
Crude protein, % 15.1 5.1 8.3 24.3 14
17.2
2.5
14.5 19.6
3 13.3
Crude fiber, % 33.1 5.2 22.6 40.6 10
29.6 21.5 37.6
2 40.3
NDF, % 53.9 2.9 50.6 55.8 3
48.1
1
ADF, % 30.7 9.4 17.3 39.7 5
38.2
1
Lignin, % 7.8 1.4 6.1 9.0 4
Ether extract, % 2.4 0.6 1.5 3.3 10 3.3 2.6 3.9 2 2.5
Ash, % 9.3 2.6 5.6 13.6 11 8.1 0.3 7.8 8.3 3 9.3
Gross Energy, MJ/kg 18.5 6.3 6.2 26.4 *
18.7
* 18.7
Calcium, g/kg 12.3 0.6 1.9 4.1 11
10.9
7.0
14.8
2 18.3
Phosphorus, g/kg 2.4 7.4 1.9 27.7 12 1.1 1 1.0
Potassium, g/kg 13.2 0.6 2.6 4.2 8
14.7 10.0 19.3
Sodium, g/kg 0.3 1 0.6
Magnesium, g/kg 3.3 8 2.7 2.5 3.0
Manganese, Mg/kg 438
Zinc, Mg/kg 27
Copper, Mg/kg 10
Iron, Mg/kg 162
Arginine, % 4.0 1
Cystine, % 1.1 1
Glycine, % 4.5 1
Histidine, % 2.7 1
Isoleucine, % 3.9 1
Leucine, % 6.7 1
Lysine, % 4.4 1
Methionine, % 1.8 1
Phenylalanine, % 4.2 1
Threonine, % 4.2 1
Tryptophan, % 2.4 1
Tyrosine, % 3.3 1
Valine, % 4.5 1
OM digestibility,
ruminants, % 62.0 *
65.6
* 55.1
Energy digestibility,
ruminants, % 59.3 *
62.8
* 51.7
DE ruminants, MJ/kg 11.0 *
11.8
* 9.7
ME ruminants, MJ/kg 8.7 * 9.4 * 7.7
Nitrogen digestibility,
ruminants, % 85.0 1 65.6
A
Avg: average or predicted value,
B
SD: standard deviation,
C
Min: minimum value,
D
Max: maximum value,
ENb: number of values (samples) used, * Indicates that the average value was obtained using an equation.
Table 4.
Kudzu’s (P. montana) (kudzu aerial part (fresh), kudzu hay, and kudzu leaves (fresh))
nutrient composition compared with nutrient requirements of various domestic ruminant species (DM
basis) [19,20,22–33].
Composition ARequirements B
Item
Kudzu,
Aerial Part,
Fresh
Kudzu
Leaves,
Fresh
Kudzu
Hay
Pregnant
Replacement
Heifers
Lactating
Beef
Cows
Lactating
Dairy
Cows
Lactating
Ewes
Lactating
Does
TDN, % 59.30 55.99 51.70 53.01 51.57 70.20 60.43 59.50
CP, % 15.10 17.20 13.30 7.82 7.56 15.60 11.20 14.50
ME, MJ/kg 8.70 9.40 7.70 4.61 7.93 11.20 9.15
Ca, % 1.23 1.09 1.83 0.25 0.21 0.56 0.32 0.55
P, % 0.24 0.11 0.10 0.19 0.14 0.36 0.23 0.33
A
Average or predicted values,
B
Average across a variety of months since conception, months since calving, milk
yield, and stages of lactation.
Agriculture 2019,9, 220 6 of 15
In situ dry matter rumen degradation data provided by Corley et al. [
20
] separates kudzu into
soluble, degradable, and indigestible fractions between leaf/stem and tuber (roots) (Table 5).
Table 5.
In situ dry matter disappearance (DMD) and
in vitro
dry matter digestion (IVDMD) of kudzu
(P. montana) plant parts [20].
In Situ DMD, % IVDMD, %
Soluble Fraction Degradable Fraction Indigestible Fraction
Leaf and Stem 29.1 48.6 22.4 64.8 (Leaf)
73.7 (Stem)
Tuber 38.1 31.2 30.7 59.9
Legumes, such as kudzu and alfalfa, have a high rate of degradability due to a low concentration
of water-soluble carbohydrates [
34
]. Corley et al. [
20
] provided data that kudzu contains 17.5% CP in
leaves, similar to alfalfa at 18.7% CP (Table 2). Kudzu leaves contain a high concentration of CP, making
it a potential feed for growing ruminants [
20
]. The stem and tuber portions of kudzu do not have the
same potential in regard to providing optimal nutrients to the ruminant. Moreover, kudzu leaves
contain a significantly higher CP level than stems; however, kudzu leaves contain significantly lower
ADF (a common predictor of energy level in forages) [
22
]. Corley et al. [
20
] found a 5–7% lower CP
level in stems than leaves (Table 6). In addition, Corley et al. [
20
] observed that kudzu tuber contained
8.6% CP (Table 6). Zhao et al. [
35
] reported that kudzu roots contain a range of CP from 3.18–4.58%. As
growth time of kudzu root increases, CP tends to decrease to levels below 4.58%. In contrast, lipid
content of kudzu root increases with an extended growth period to levels above 32.2 g/kg [35].
Table 6. Nutritive values of kudzu (P. montana) plant parts [20].
Parameters Leaf Stem Tuber
Crude protein, (CP), % 17.5 10.3 8.6
Neutral detergent fiber, (NDF), % 48.1 73.7 39.8
Acid detergent fiber, (ADF), % 38.2 44.0 53.3
Ca, % 0.7 0.1 0.4
K, % 1.0 1.0 0.3
Mg, % 0.3 <0.1 0.1
Fe, mg/kg 162.3 156.6 3600.0
Additionally, Gulizia et al. [
36
] observed that kudzu from two different growing seasons contained
a higher concentration of nutrients than in previous reports (Table 7). This study also observed that
early season kudzu dry matter degradability was 84%, and late season kudzu degradability was 79%
over a 72 h incubation period. Gulizia et al. [
36
] concluded that kudzu (regardless of growing season)
was highly degradable over a 72 h incubation period in ruminants, and has potential as a feedstock.
In comparison, alfalfa is a high-quality forage characterized by high digestibility and swift ruminal
degradation [
37
]. Alfalfa and kudzu leaf and stem have similar in situ rumen degradation, with alfalfa
having an average rate of 73.35% and kudzu having 78% maximum degradability [
20
,
38
]. Alfalfa
leaf and stem have an average soluble fraction of 34.8% and an average degradable fraction of 38.6%,
whereas kudzu leaf and stem are 29.1% and 48.6%, respectively (Table). This data can potentially
predict that alfalfa leaf and stem contain more starch, sugars, and protein than kudzu leaf and stem,
but less concentrations of cellulose and hemicellulose. Coblentz et al. [
38
] allowed alfalfa to ferment in
the rumen for 96 hours, whereas Corley et al. [
20
] only allowed 24 hours of fermentation for kudzu. In
situ dry matter disappearance for whole plant alfalfa was found to be 76.6, 79.6, 79.2, and 81.91% in
four studies [
38
–
41
]. It is common for alfalfa to have high nitrogen levels that are highly degradable.
High degradability of nitrogen in alfalfa can lead to poor utilization of available nitrogen in lactating
dairy cows [38].
Agriculture 2019,9, 220 7 of 15
Table 7.
Near infrared reflectance spectroscopy (NIR) chemical composition of early (ES) and late
season (LS) kudzu (P. montana) leaves incubated in experimental fistulated bovine during in situ rumen
degradation study periods (DM basis) [36].
Analysis ES LS
DM, % 88.2 88.5
CP, % 30.5 26.7
Available protein, % 28.9 24.9
ADF, % 18.4 26.2
NDF, % 27.3 45.7
Lignin, % 4.40 6.3
Net Energy (maintenance), Mcal/kg 1.80 1.23
Net Energy (growth), Mcal/kg 1.19 0.68
Ca, % 1.54 1.74
P, % 0.43 0.28
Mg, % 0.35 0.39
K, % 2.05 2.05
S, % 0.34 0.39
Cl, % 0.45 0.55
Lysine, % 1.55 1.25
Methionine, % 0.48 0.40
6. Anti-Quality and Anti-Nutritional Factors of Kudzu
Kudzu contains a variety of secondary metabolites. Table 8summarizes a qualitative analysis
of some common secondary metabolites found in tropical kudzu. Data is limited on quantitative
analysis of many secondary metabolites in kudzu leaf and vine. These secondary metabolites can act
as anti-nutritional or anti-qualitative factors [
42
,
43
]. Legumes, such as kudzu, are beneficial in nitrogen
fixation and improvement of animal diets. Secondary metabolites in kudzu, however, can interfere
with nutrient intake, absorption, and utilization [42,43].
Table 8.
Summary of qualitative tests from phytochemical screening and analysis of kudzu
(Pueraria phaseoloides) [43].
Secondary Metabolites Tropical Kudzu
Organic acids +++
Reducing Sugars +++
Saponins +++
Tannins +++
Steroids and triterpenoids ++
Saccharides ++
Alkaloids +
Depsides and depsidones +
Coumarin in by-products +
Flavonoids +
Cardiac glycosides +
Catechins -
Lactones -
Purines -
Quinones -
(+++) large presence; (++) medium presence; (+) small presence; (-) absence or inconclusive result of
secondary metabolites.
There are both toxic (i.e., alkaloids, saponins, isoflavones, etc.) and nontoxic (i.e., tannins, cutin,
biogenic silica, etc.) secondary metabolites in plant materials [
43
,
44
]. Alkaloids, cyanogenic glycosides,
toxic amino acids, saponins, and isoflavones are toxic compounds present in low concentrations [
43
,
44
].
These compounds at concentrations less than 20 g/kg can have negative effects when absorbed by an
Agriculture 2019,9, 220 8 of 15
animal, including neurological problems, reproductive failure, gangrene, and potential fatalities [
43
,
44
].
Lignin, tannin, cutin, biogenic silica, and volatile terpenoids make up the non-toxic compounds present
in high concentrations. These compounds at concentrations greater than 20 g/kg can result in decreased
digestibility and palatability [43,44].
Saponins are in high concentrations in tropical kudzu (Pueraria phaseoloides), causing tympanism
(accumulation of gas), reduced rumen microbial fermentation, and hepatic photosensitivity [
43
].
Saponins also create stable foam in water and impart a bitter flavor to forages, thus decreasing the
likelihood of intake by the animal [
43
]. Saponins are major anti-nutritional and anti-qualitative
factors, but tannins are the primary negative factor in legumes [
43
]. There are two types of tannins,
hydrolysable and condensed varieties, with the latter being found in legumes, sorghum grains, and
tree leaves. Tannins contain a large amount of phenolic hydroxyl groups, allowing them to create links
with proteins and other molecules [
45
]. A main concern with tannins in feedstuffs is their negative
effects on the ruminant digestive system through protein interactions [
45
]. Tannins will affect the
nutritive value of plant dry matter, reducing the palatability by precipitating salivary proteins and
nutrient digestibility by diminishing the permeability of the rumen wall through interactions with the
outer cellular layer of the digestive tract. Tannins consumed at >50 g/kg of dry matter concentration
will cause ruminants to reject feedstuffs, while consumption <50 g/kg seems to not affect voluntary
feed intake [
46
–
49
]. Digestive enzyme activity may also decrease from tannins’ ability as a potent
inhibitor. Tannins have the potential to cause negative effects to an animal, including impaired ruminal
digestion; low milk yield; toxic degenerative changes in the intestine, liver, spleen, and kidney; and
constipation [
45
]. Both saponins and tannins cause negative effects, but the positive effects these
secondary metabolites can have, including diminished ruminal methane production, is still being
explored [43,50].
Organic acids and reducing sugars are the remaining secondary metabolites that have large
concentrations in kudzu. Organic acids can bring about precipitation of calcium ions in the blood,
leading to muscle weakness, nephritis, kidney stones, gastrointestinal irritation, and hypocalcemia
syndrome in grazing ruminants and horses [
43
]. In large concentrations, reducing sugars can be
problematic for equines [
43
,
51
]. Equines fed a high concentrate diet will produce excess lactic acid,
resulting in water retention and decreased pH values in the lumen of the digestive tract [
43
,
51
]. This
risks the possibility of digestive disorders, including osmotic diarrhea and colic [
43
,
51
]. Additional
secondary metabolites, such as coumarin by-products, depsides and depsidones, alkaloids, steroids,
triterpenoids, flavonoids, and cardiac glycosides can also cause negative effects [
43
]. Lastly, kudzu
contains phenolic compounds that can have allelopathic advantages [
52
,
53
]. Kudzu leaves and roots
contain 2–3% (DM) phenolic compounds [
52
,
53
]. Kudzu growing soils contain approximately 50 times
more phenolics than soils devoid of kudzu [52,53].
7. Biological Control of Kudzu Using Animals
Plant populations are controlled naturally by their environment and by natural enemies. Invasive
species are unique in disrupting an ecosystem to which it does not belong due to a lack of natural
control [
54
]. Invasive species are the second largest cause of biodiversity (total variability within and
among species of all plant organisms and their habitats) loss, behind habitat destruction [
55
]. The
degree of invasiveness may increase with a lack of natural competitors [
55
,
56
]. Plant species that
are established in an environment outside of its natural habitat may be less regulated by the native
herbivores in the area, thus resulting in the rapid growth of an invasive plant species [
55
,
56
]. This leads
to unwanted imbalances in an ecosystem that have potential to harm native species. These invasive
plant species can be controlled by chemical or biological methods. Biological control is a method by
which one organism is used to control another and can be used to restore ecosystem balance [
56
].
Biological control of problematic species using animals was recorded as early as 9,500 years ago when
cats were domesticated to control rodents [
56
]. Animals used in biological control of invasive plant
species can range from insects to ruminants. Therefore, land owners can manage livestock to use
Agriculture 2019,9, 220 9 of 15
invasive plant species as diet supplements and not only enhance animal production, but also slowly
diminish the infestation of the invasive species.
Kudzu (P. montana) must undergo constant application of some control method to yield results
in lowering its occurrence. Efforts to successfully control this plant is heavily influenced by timed
treatments within its life cycle [
6
]. Biological control using grazers and browsers can be an effective and
cost-efficient method, but it is a slow process [
6
]. Elimination of kudzu is possible by frequent defoliation
by animals over several years. By over defoliating and reducing photosynthetic carbon, hydrogen, and
oxygen (CHO), kudzu will halt its metabolic processes and regrowth will be prevented [
2
]. Defoliation
during the fall will reduce the amount of resources roots receive for survival through the winter, thus
accelerating the progression of eradication [5].
Kudzu (P. montana) can be eliminated using cattle to over graze it at 80% consumption of the
vegetative growth for 3–4 years [
2
,
6
]. In contrast, tropical kudzu (Pueraria phaseoloides) could be
efficiently controlled using cattle on a rotational grazing system in less than 2 years [
57
]. However,
for tropical kudzu to be eradicated in 2 years, the soil should be compact and drain poorly [
57
].
Vines which these animals cannot reach may be cut and fed to ensure that defoliation is effective.
Remaining plant material after those 3–4 years can be spot treated with recommended herbicides [
6
].
Furthermore, continuous grazing and browsing of infested areas for approximately 2 months during
kudzu’s growing season (May-October in the U.S.) can be effective in its eradication. Older infestations
become increasingly hard to eradicate. Kudzu over 10 years old will be minimally affected by over
grazing and over browsing, so herbicide application may be necessary [
6
–
8
]. However, kudzu’s hardy
nature tends to make application of herbicides difficult due to the stockpile of starch in its tap root [
58
].
8. Use of Kudzu as a Feedstock
Grazers (e.g., cattle and sheep) and browsers (e.g., goats and deer) will consume kudzu (P. montana)
when available, but it is easily overbrowsed or overgrazed [
4
]. Kudzu is known to produce a forage of
high quality that contains a crude protein (CP) concentration of 15% or higher and a total digestible
nutrient (TDN) value of over 60%, but the use of this plant as a feedstuffhas limitations. Kudzu grows
rapidly, but it produces a low forage yield of 2–4 tons of dry matter per acre per year [
2
]. Pairing
low forage yield with a vine-like growth habit makes harvesting problematic. During dry periods,
producers can harvest kudzu annually or biennially, as it retains moisture for growth deep within the
roots [
2
]. Grazers and browsers can be enclosed on a plot of kudzu to control its growth, while also
receiving a high quality source of nutrients that potentially results in increased animal performance.
Lynd and Ansman [
59
], Miller and Edwards [
60
], and Rhoden et al. [
61
] reported that heavy grazing
kudzu for 3–4 growing seasons with cattle, swine, horses, sheep, or goats in August and September
could prove to be effective at starving and preventing growth of kudzu. Though data is limited, there
are examples of kudzu’s use as a feedstock.
Tropical kudzu contains a high nutrient composition as shown in Table 9. Overall, P. montana and
Pueraria phaseoloides tend to be similar in nutritive value. Previous research utilizing tropical kudzu
determined that it is palatable and contains an adequate amount of CP for ruminants. Monteiro et al. [
62
]
observed that dairy cows fed a diet that consisted of tropical kudzu supplemented with sorghum grain
could support a milk yield of 8.1 kg milk/day.
Agriculture 2019,9, 220 10 of 15
Table 9.
Chemical composition and nutritional value of tropical kudzu (Pueraria phaseoloides), aerial
part, fresh (DM basis) [26,63–71].
Item Avg ASD BMin CMax DNb E
Dry matter, % as fed 19.0 4.0 14.0 32.7 33
Crude protein, % 19.3 3.3 13.1 25.8 48
Crude fiber, % 33.0 3.9 26.7 40.2 42
NDF, % 49.4 46.3 51.9 2 *
ADF, % 38.2 5.1 28.4 38.5 3 *
Lignin, % 7.1 1.6 5.4 8.5 3
Ether extract, % 2.2 0.7 1.0 3.9 38
Ash, % 8.7 1.7 5.3 11.3 46
Gross Energy, MJ/kg 18.9 *
Calcium, g/kg 9.6 2.2 5.4 14.5 42
Phosphorus, g/kg 2.7 0.7 1.5 4.0 41
Potassium, g/kg 23.6 7.2 10.2 36.5 39
Sodium, g/kg 0.1 0.1 0.1 0.2 3
Magnesium, g/kg 3.0 0.5 2.1 4.1 38
Manganese, mg/kg 98 43 153 2
Zinc, mg/kg 40 39 42 2
Copper, mg/kg 12 12 13 2
Iron, mg/kg 206 1
Arginine, % 4.1 4.0 4.3 2
Cystine, % 1.1 1
Histidine, % 1.8 1.7 1.9 2
Isoleucine, % 3.9 3.5 4.4 2
Leucine, % 6.4 5.8 7.0 2
Lysine, % 3.3 3.2 3.3 2
Methionine, % 1.8 1
Phenylalanine, % 4.5 4.4 4.6 2
Threonine, % 4.4 4.3 4.4 2
Tryptophan, % 1.2 1
Tyrosine, % 3.4 1
Valine, % 4.6 4.3 4.9 2
OM digestibility, ruminants, % 62.2 *
Energy digestibility, ruminants, % 59.4 *
OF ruminants, MJ/kg 11.2 *
ME ruminants, MJ/kg 8.8 *
Nitrogen digestibility, ruminants, % 80.0 1
A
Avg: average or predicted value,
B
SD: standard deviation,
C
Min: minimum value,
D
Max: maximum value,
ENb: number of values (samples) used, * Indicates that the average value was obtained using an equation.
In 1945, tropical kudzu (Pueraria phaseoloides) was introduced to a herd of Guernsey cows. Initially,
few cows ate kudzu, but within days cows consumed it regularly [
63
]. This experiment was conducted
when common pasture crops did not produce a sufficient forage. For one continuously grazing cow
during the dry season, an estimated one acre of tropical kudzu was needed. Telford and Childers [
63
]
determined that tropical kudzu should only be grazed once during the dry season. To use kudzu for
grazing, it should not be grazed to the ground to preserve quality and regrowth ability. Tropical kudzu
had an estimated 11,000–18,150 kg of forage production per year during these experiments. Successful
grazing was also established using oxen and goats, and an adapted use for other livestock and poultry.
Kudzu has been used as a source of feed for a variety of research animals. Bhatt and Sharma [
23
]
fed fresh kudzu-vine (Puereria thunbergiana) ad libitum to experimental Angora rabbits. In the Malagasy
Republic, both Puereria thunbergiana and Pueraria phaseoloides were successfully grown as a high protein
feedstuff[
26
,
58
]. Using kudzu as an alternative feed source has been successful due to its palatability
and positive results have been shown when fed to beef and dairy cattle [
22
]. Cows utilizing kudzu
as a feedstock produced milk with no color or flavor differences [
22
]. Piper [
72
] and Shurtleffand
Aoyagi [
73
] observed that when given a choice, cattle, hogs, chickens, goats, sheep, horses, and rabbits
Agriculture 2019,9, 220 11 of 15
preferred kudzu over grasses or commercial hay. Research conducted in Alabama on Angora goats
observed that kudzu populations were effectively controlled with 27 animals/hm
2
[
52
,
74
]. Additionally,
kudzu is ideal as an emergency feed source when common sources are low [
22
]. Polk and Gieger [
75
]
demonstrated that when alfalfa became limited, kudzu meal at 9% of their diet could be substituted in
chick rations. Kudzu and various grasses (e.g., par
á
grass) can form a desirable combination (when
grazing) to increase protein content and reduce the need for commercial feeds [
63
]. Poultry have been
observed to graze kudzu and consume kudzu seeds; however, there have been no reports on potentially
using poultry as a way to biologically control kudzu [
52
,
59
]. Nworgu and Egbunike [
76
] researched
potential growth performance effects of feeding tropical kudzu (Pueraria phaseoloides) leaf meal to
broilers. They concluded that, although the tropical kudzu leaf meal was rich in nutrients, tropical
kudzu leaf meal should not be included in broiler diets as it led to poor growth rates. However, there
are a growing number of researchers that are assessing leaf meals as an alternative protein supplement
in modern poultry diets [
77
,
78
]. Thus, further research should assess the potential viability of kudzu’s
use as a protein supplement in countries that have limited access to modern protein supplements (e.g.,
soybean meal). This would indicate that kudzu can be of real relevance for non-herbivore species such
as poultry.
9. Summary and Applications
Kudzu’s high quality nutrient composition, degradability, and palatability suggest that it can be
a valuable potential feedstock for livestock species. Its widespread distribution and rapid growth
rate make it ideally suited as an economical feedstock, particularly in regions of scarce feedstuff
availability. Producers must recognize, however, the challenges of managing kudzu for grazing and
browsing species. With kudzu’s unique growth habits as a climbing vine, containing livestock species
in areas of high kudzu mass can be challenging and may require considerable ingenuity. Furthermore,
efforts to prevent overgrazing/overbrowsing should be highlighted. Kudzu leaf volume can be quickly
depleted, effectively limiting kudzu’s use as a feedstock. Overgrazing/overbrowsing would be a
preferred approach if the producer was attempting to control kudzu with livestock, but not if using as
a sustainable source of nutrients.
This review has attempted to coalesce an array of information on kudzu into a comprehensive
overview of its characteristics and potential significance as a feedstufffor a variety of domestic animal
species. Much information, however, remains to be elicited on kudzu. Kudzu is often viewed in
a negative light as an invasive weed species. Kudzu is, however, much more than that and holds
significant potential as a feedstock in the United States and other countries where nutrient availability
for production animals is limited.
Author Contributions:
Conceptualization, J.P.G. and K.M.D.; Methodology, J.P.G. and K.M.D.; Validation,
J.P.G. and K.M.D.; Formal analysis, J.P.G. and K.M.D.; Investigation, J.P.G. and K.M.D.; Writing—original draft
preparation, J.P.G.; Writing—review and editing, J.P.G. and K.M.D.; Visualization, J.P.G. and K.M.D.
Funding: This research received no external funding.
Conflicts of Interest: The authors declare no conflict of interest.
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