White
Horsenettle (Solanum elaeagnifolium)
From
the Washington State Noxious Weed Control Board
Send
questions or comments about this page to Dana Coggon at the Washington
State Noxious Weed Control Board (360)902-2082
Silverleaf
nightshade aka. White Horsenettle is a branched, deep-rooted,
perennial herb, 1 to 4 feet
tall. Slender, yellow spines occur on the stems or leaf ribs of
the plant. The lance-shaped leaves are 1 to 4 inches long by
1 inch
wide, with wavy margins; they are covered with short, silvery-white,
star-shaped hairs that give the plant a dusky or silvery-gray
color.
The blue, violet or rarely white flowers have 5 fused petals, ¾
inch across, with bright yellow stamens. Flowers grow on stalks
in clusters or singly at the end of stems or branches. The fruits
are yellow to brownish, juicy berries, ½ inch in diameter.
Seeds are flat, brown and 1/10 to 1/5 inch long (Boyd et al. 1984;
Gunn and Gaffney 1974; Roche 1991; Rutherford 1978).
The
Solanaceae family includes food plants, poisonous and medicinal
plants, ornamentals, and several noxious weeds (Boyd et al. 1984).
Economic
Importance
Beneficial:
Silverleaf nightshade is rich in solasodine, a chemical used in
the manufacture of steroidal hormones. A protein-digesting hormone
resembling papain is present in its fruits. Pima Indians added crushed
berries to milk when making cheese. The Kiowa Tribe combined silverleaf
nightshade seeds with brain tissue and used it for tanning hides
(Boyd et al 1984).
Detrimental
Silverleaf
nightshade lowers crop yield through competition (Boyd and Murray
1982b). The plant grows in the early spring due to
food
reserves in its well-developed root system. The roots grow deeper
than those of associated crops. These traits may give it an
advantage over agricultural species, including wheat, alfalfa,
cotton,
peanuts,
and grain sorghum (Boyd et al. 1984; Roche 1991). In addition,
the plant’s spiny leaves and coarse stems may lower the quality
of hay taken from infested fields (Boyd et al. 1984).
The
species is also toxic to livestock. Silverleaf nightshade contains
toxic alkaloids that combine with sugars to produce glycoalkaloids
that irritate the gastrointestinal tract; within the tract, these
compounds may be hydrolyzed to release alkalids or alkamines that
are nerve toxins (Boyd et al. 1984). Cattle that consume 0.1% to
0.3% of their body weight in ripe berries display moderate poisoning
symptoms, which may include: rapid, labored breathing; salivation
and slobbering; nasal discharge; yellow discoloration of the skin
in light-colored animals; weakness and lack or coordination; trembling
of muscles in back legs; anemia; and increased heart rate (Buck
et al. 1960). Sheep are more resistant to the toxins and goats are
unaffected (Boyd et al. 1984).
Silverleaf
nightshade can also harbor plant pests, such as lygus bugs, Colorado
potato beetle, and leafspot (Boyd et al. 1984; Roche 1991).
Geographical
Distribution
Solanum
elaeagnifolium is native to the Americas, although it is unclear
whether it originated in North America
or
South America; Spanish or Portuguese colonists may have moved
the species from North America to South America or vice versa.
However,
the most likely center of geographic origin is the southwestern
U.S. or northern Mexico (Boyd 1984). Interest in the plant
increased in the 1970’s as silverleaf nightshade spread outside its
native range. It is known from Australia, Egypt, Greece, India,
Israel, Zimbabwe, Sicily, South Africa, Morocco and Spain (Boyd
et al. 1984; Bouhache and Tanji 1985). It is a listed noxious
weed
in 21 states (Roche 1991). In the Pacific Northwest, the plant
has been introduced to Umatilla County, Oregon; Idaho County,
Idaho;
and Asotin and Walla Walla counties, Washington (Roche 1991; Washington
State Noxious Weed Control Board, unpublished data).
Habitat
Silverleaf
nightshade is adapted to semi-arid regions with 12 to
23 inches of annual rainfall. The plant typically occurs on coarse-textured,
sandy soils (Molnar and McKenzie 1976 cited in Boyd et al.
1984). In its native range, silverleaf nightshade is a problem
in areas
where the vegetation has been removed, such as roadsides, construction
sites, livestock feeding and watering areas, and cultivated
fields. It is considered a problem in cereal grain, alfalfa,
grain sorghum
and cotton. In California, where the plant was introduced,
it occurs
in orchards, agronomic and vegetable crops, roadsides, pastures,
and vacant lots (Roche 1991).
History
There
is no record of how silverleaf nightshade was introduced to
Washington. In California, the plant became established along
railroad tracks after it was swept from railcars with bedding
material
(Roche
1991).
Growth
and Development
Silverleaf
nightshade is a summer-growing perennial
plant, with an extensive root system. Roots can grow very deep
(6 to 10 feet) and extend horizontally to produce shoots 6
feet away
from the parent plant (Roche 1991; Davis et al. 1945). Shoots
start to emerge from established plants as the soil warms in
late
March
to early April. Plants may begin to flower in early May (Cooley
and Smith 1971). Ripe fruits may be present in June, and some
seeds
are viable the season they are produced. Seedlings may appear
in August and September in flooded areas. Plants die back in
winter
and reappear from roots in the spring (Davis et al. 1945).
Reproduction
Solanum
elaeagnifolium can spread by seed, rhizomes, and/or root
fragments (Boyd and Murray 1982b). Flowers are cross-pollinated
by insects (Buchmann and Cane 1989). Individual berries produce
24 to 149 seeds (Boyd and Murray 1982b), which can add up to 5
million to 100 million seeds per acre (Cooley and Smith 1971).
Seeds may
be dispersed by wind, water, machinery, agricultural produce
or animal feces; studies indicate that 10% of seed is still viable
after passing through sheep. Dried plants may also blow like tumbleweeds,
spreading seed along the way (Boyd et al. 1984).
Seeds
require fluctuating temperatures to germinate. Boyd and Murray (1982b)
obtained a maximum germination rate of 57% when they germinated
seeds at 20° C for 16 hours and 30° C for 8 hours; light
and dark had no effect. They also found that a pH of 6 or 7 was
optimal for germination. Other work indicates that immersing seeds
in running water for relatively long periods may improve germination
rates (Rutherford 1978).
Plants
may also spread by rhizomes or root fragments. Local distribution
of the plant’s vegetative propagules is usually the result
of tillage and the creeping nature of the rhizomes (Boyd et al.
1984). Rhizomes may extend 6 feet from the parent plant (Molnar
and McKenzie 1976 cited in Boyd et al. 1984). Root fragments as
small as 0.4 inches can regenerate (Richardson and McKenzie 1981),
and sections of taproot may maintain their viability for up to 15
months (Molar and McKenzie 1976 cited in Boyd et al. 1984).
Response
to Mechanical Control Methods
Plants
can regrow after being clipped (Cooley and Smith 1971). Removing
plant tops by mowing results
in
a loss of apical dominance that causes multiple shoots to re-sprout
(Boyd and Murray 1982b). Removing above-ground parts every
2 weeks
can prevent seed production (Roche 1991).
Response
to Cultural Control Methods
Cultivation
will not readily kill silverleaf
nightshade (Richardson and McKenzie 1981). Only frequent, thorough
cultivation can be effective. Reduced tillage agriculture produces
longer root fragments, which result in more shoots and faster
growth.
As a result, there is more interference with crop production,
and it is more difficult to control the weed (Boyd and Murray
1982a).
Shade
from crop canopies can be an effective control tool. However, silverleaf
nightshade must be restrained in some other way until crop canopy
formation is complete. Otherwise, the weed will mature first and
be less impacted by shade. Shade levels between 63% and 92% are
needed to prevent seed production. Shade from crop canopies decrease
silverleaf nightshade’s photosynthetic rate, which may make
it a less vigorous competitor (Boyd and Murray 1982a). In Arizona,
eradication was achieved in 3 years by hoeing silverleaf nightshade
to the ground until grain sorghum formed a dense canopy (Roche 1991).
Response
to Herbicides
The
plant is difficult to control with herbicides because the root
system is widespread and connects to adjacent
above-ground
growth (Richardson 1979). Studies in California and Greece indicated
that glyphosate and picloram provide consistent control of
this weed (Eleftherohorinos et al. 1993). California growers
have
used
soil fumigation to eradicate small infestations (Roche 1991).
For specific herbicide recommendations, refer to the Pacific
Northwest
Weed Control Handbook.
Biocontrol
Potentials
In
Arizona, New Mexico, and Texas, part of the plant’s
native range, 116 insects were collected on silverleaf nightshade.
However, it is still weedy in these areas (Goeden 1971). Several
biocontrol agents have been investigated. The most promising organism
is a foliar nematode, Orrina phyllobia, which causes leaf and stem
galling (Roche 1991).
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