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thesis entitled

EVALUATION OF NIGHTSHADE (Solanum spp.) AND
GROUNDCHERRY SPECIES (Physalis spp.) RESPONSE TO

HERBICIDES

presented by

VIJAIKUMAR PANDIAN

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EVALUATION OF NIGHTSHADE (Solanum spp.) AND GROUNDCHERRY
SPECIES (Physalis spp.) RESPONSE TO HERBICIDES
By

Vijaikumar Pandian

A THESIS
Submitted to
Michigan State University
in partial fulfillment of the requirements for the degree of
MASTER OF SCIENCE
Department of Horticulture

2005

ABSTRACT
EVALUATION OF NIGHTSHADE (Solanum spp.) AND GROUNDCHERRY
SPECIES (Physalis spp.) RESPONSE TO HERBICIDES
By
Vijaikumar Pandian
Solanaceous weeds such as eastern black nightshade (Solanum ptycanthum), hairy
nightshade (Solanum sarrachoides), and horsenettle (Solanum carolinense) are serious
weeds in tomato production in Michigan. Groundcherries, such as clammy groundcherry
(Physalis heterophylla) and smooth groundcherry (Physalis subglabrata) are usually less
troublesome weeds in tomatoes. Studies were conducted in the ﬁeld and greenhouse to
determine nightshade and groundcherry response to herbicides. Germination studies were
conducted in growth chambers to determine the inﬂuence of temperature on germination
of nightshade and groundcherry populations collected in Michigan. Post-transplant
application of S-metalochlor (1.8 kg/ha), and dimethenamid-P (1.09 kg/ha) and pre-
transplant application of ﬂumioxazin (0.05 kg/ha), and oxyﬂuorfen (0.28 kg/ha) in the
ﬁeld gave 95% control of eastern black nightshade with no tomato injury. Postemergence
application of pyridate (1.01 kg/ha) gave 60% control of eastern black nightshade in ﬁeld
with no crop injury. There was variation in dose response Of eastern black nightshade
populations to sulfosulfuron, halosulfuron, and metribuzin in the greenhouse. There was
variation in dose response of hairy nightshade populations to halosulfuron. Eastern black
nightshade had a higher germination rate than horsenettle, hairy nightshade, smooth and
clammy groundcherries at 28/20 °C. An eastern black nightshade population from Oceana

County germinated at a wider range of temperatures from 28/200 C to 15/ 10 ° C.

ACKNOWLEDGMENTS

Om. First of all, I Offer my prayer to Lord Ganesha “the remover of all Obstacles”. I
praise the feet of Lord Shiva, for his proffering grace throughout my life.

This dissertation would not have been possible without the assistance Of the
committee members, Dr. Bernard H. Zandstra, Dr. Karen Renner and Dr. Mathieu
Ngouajio. Their guidance and support in my professional life are valuable. I would like to
thank my major professor, Dr. Bernard H. Zandstra for Offering the opportunity to pursue
my Master degree program in Horticulture at Michigan State University. I appreciate his
great patience. I would like to extend my heartfelt thanks to all my co-workers for their
selﬂess help to make this research work successful.

Above all, I would like to thank my parents, my brother, my sister and my teacher

- Thiru Sankaran, for their endless love, support and affection upon me.

TABLE OF CONTENTS

LIST OF TABLES .......................................................................................................... vii

LIST OF FIGURES .......................................................................................................... x
CHAPTER 1

LITERATURE REVIEW ................................................................................................. 1

INTRODUCTION ................................................................................................... 1

TAXONOMIC CHARACTERISTICS .................................................................... 2

HISTORY ................................................................................................................ 3

SEED PRODUCTION AND DISPERSAL CHARACTERISTICS ....................... 3

GERMINATION CHARACTERISTICS ................................................................ 4

GROWTH AND PHYSIOLOGICAL CHARACTERISTICS ................................ 5

INTERFERENCE IN TOMATO GROWTH AND DEVELOPMENT .................. 6

ALTERNATE HOST TO PEST AND DISEASES ................................................. 7

DIFFICULT TO CONTROL IN TOMATO ............................................................ 8

LITERATURE CITED .......................................................................................... 10
CHAPTER 2

POTENTIAL HERBICIDES TO CONTROL EASTERN BLACK NIGHTSHADE

(Solan um ptycanth um) IN TOMATO (L ycopersicon esculentum) ............................... 15

ABSTRACT ........................................................................................................... 15

INTRODUCTION ................................................................................................. 16

MATERIALS AND METHODS ........................................................................... 18

FIELD STUDY .......................................................................................... 18

FIELD PREPARATION ................................................................ l8

TRANSPLANTIN G ...................................................................... 1 8

APPLICATION METHODS ......................................................... 19

VISUAL RATING ......................................................................... 19

YIELD ............................................................................................ 19

STATISTICAL ANALYSIS ......................................................... 20

GREENHOUSE STUDY ........................................................................... 20

PRE TREATMENT OF NIGHTSHADE SEEDS ......................... 20

SOWING AND TRANSPLANTING ............................................ 21

TREATMENT APPLICATION .................................................... 21

VISUAL RATING AND HARVESTING OF BIOMASS ............ 22

STATISTICAL ANALYSIS ......................................................... 22

RESULTS AND DISCUSSION .......................................................... 23

FIELD STUDY ..................................................................... 23

PRE - TRANSPLANT HERBICIDE .................................. 23

POST - TRANSPLANT HERBICIDE ................................ 24

POSTEMERGENCE HERBICIDE ...................................... 25

POST — DIRECTED HERBICIDE ...................................... 26

GREENHOUSE STUDY ........................................................................... 26

DOSE RESPONSE OF SULFOSULFURON.................. .......26

DOSE RESPONSE OF SULFENTRAZONE................. .........27
LITERATURE CITED ...................................................................... 28

CHAPTER 3
VARIATION IN HERBICIDE RESPONSE AMONG NIGHTSHADE (Solanum
spp) AND GROUNDCHERRY (Physalis spp) SPECIES AND WITHIN

NIGHTSHADE POPULATION ................................................................. 40
ABSTRACT ................................................................................... 40
INTRODUCTION ............................................................................ 41
MATERIALS AND METHODS ............................................................ 43

COLLECTION OF BERRIES ..................................................... 43
PROCESSING OF BERRIES ..................................................... 43
PRE TREATMENT OF SEEDS ................................................... 43
SOWING AND TRANSPLANTING ............................................. 44
TREATMENTS AND APPLICATION .......................................... 44
VISUAL RATING AND BIOMASS MEASUREMENTS. . 44
STATISTICAL ANALYSIS ...................................................... 45
RESULTS AND DISCUSSION ............................................................ 46
RESPONSE OF EASTERN BLACK NIGHTSHADE POPULATIONS
TO HERBICIDES ................................................................... 46
RESPONSE OF HAIRY NIGHTSHADE POPULATIONS TO
HERBICIDES ....................................................................... 48
RESPONSE OF HORSENETTLE POPULATIONS TO HERBICIDES .49
RESPONSE OF CLAMMY GROUNDCHERRY TO HERBICIDES ..... 50
RESPONSE OF SMOOTH GROUNDCHERRY TO HERBICIDES ...... 51
LITERATURE CITED ....................................................................... 52
CHAPTER 4

EFFECT OF TEMPERATURE ON GERMINATION OF EASTERN BLACK
NIGHTSHADE, HAIRY NIGHTSHADE, HORSENETTLE, SMOOTH

GROUNDCHERRY AND CLAMMY GROUNDCHERRY ............................ 107
ABSTRACT ............................................................................... 107
INTRODUCTION ........................................................................ 1 08
MATERIALS AND METHODS ........................................................ 111

PROCESSING OF SEEDS ..................................................... 111
VIABILITY TESTS .............................................................. 1 l 1
DESIGN OF THE EXPERIMENT ............................................. 1 12
TREATMENTS ................................................................... 112
GERMINATION COUNT ...................................................... 112
STATISTICAL ANALYSIS ................................................... 1 13
RESULTS AND DISCUSSION ......................................................... 114
GERMINATION RESPONSE OF EASTERN BLACK
NIGHTSHADE POPULATIONS ............................................. 1 14

GERMINATION RESPONSE OF HORSENETTLE

POPULATIONS ...................................................................................... 1 15
GERMINATION RESPONSE OF HAIRY NIGHTSHADE

POPULATIONS ................................................................... l 16

GERMINATION RESPONSE OF SMOOTH AND CLAMMY
GROUNDCHERRY ............................................................... 1 17
LITERATURE CITED .................................................................... 119
APPENDICES ..................................................................................... 129
LIST OF COMMON AND CHEMICAL NAMES OF HERBICIDES .......... .129

vi

LIST OF TABLES

CHAPTER 2
POTENTIAL HERBICIDES TO CONTROL EASTERN BLACK NIGHTSHADE
(Solan um ptjycanth um) IN TOMATO (Lycopersicon esculentum)

Table 1. List of treatments, application methods and the rate applied during in ﬁeld
experiments during 2002, 2003 and 2004 to evaluate the control of eastern black
nightshade in transplanted tomato .................................................................................... 30

Table 2. Pre transplant (PRT) treatments effect on eastern black nightshade (SOLPT) and
crop injury rated at six weeks after treatment in 2002, 2003 and 2004 ....................... 31

Table 3. Post transplant (POT) treatments effect on eastern black nightshade (SOLPT)
and crop injury rated at six weeks after treatment in 2002, 2003 and 2004 ................. 32

Table 4. Post emergence (POST) and post directed (PODIR) treatments effect on eastern
black nightshade (SOLPT) and crop injury rated at three weeks after treatment in 2002,
2003 and 2004 ........................................................................................ 33

Table 5. Effect of pre-transplant (PRT) treatments on tomato yield from 2002 - 2004.. ..34

Table 6. Effect Of post-transplant (POT) treatment on tomato yield from 2002- 2004 .. . .35

Table 7. Effect of post emergence (POST) and post directed (PODIR) treatments on
tomato yield from 2002-2004 .................................................................................... 36

Table 8. Polynomial quadratic equations, R2 values“, GR50 values to determine the
response of eastern black nightshade (SOLPT) and tomato to sulfosulfuron“,
sulfentrazone treatments applied during two different season ................................. 37

CHAPTER 3

VARIATION IN HERBICIDE RESPONSE AMONG NIGHTSHADE (Solanum
spp) AND GROUNDCHERRY (Physalis spp) SPECIES AND WITHIN
NIGHTSHADE POPULATIONS

Table l.Eastem black nightshade (SOLPT) populations collected from various regions in
Michigan; including location, current crop and soil properties..................... .............54

Table 2. Hairy nightshade (SOLSA) populations collected from various regions in
Michigan; including location, current crop and soil properties .............................. .55

Table 3. Horsenettle (SOLCA) populations collected from various regions in Michigan;
including location, current crop and soil properties ............................................... 56

vii

Table 4. Clammy groundcherry (PHYHE) and smooth groundcherry (PHYSU)
population collected from various regions in Michigan; including location, current crop
and soil properties ................................................................................... 57

Table 5. Treatments applied at various concentrations on eastern black nightshade. hairy
nightshade, horsenettle, smooth groundcherry and clammy groundcherry

populations ............................................................................................ 58

Table 6. Log logistic equations, R2 values and GR50 values for eastern black nightshade
(SOLPT) populations in response to metribuzin treatment ..................................... 59

Table 7. Log logistic equations, R2 values and GR50 values for eastern black nightshade
(SOLPT) populations in response to sulfentrazone treatment .................................. 60

Table 8. Log logistic equations, R2 values and GRSO values for eastern black nightshade
(SOLPT) populations in response to halosulfuron treatment .................................. 61

Table 9. Log logistic equations, R2 values and GR50 values for eastern black nightshade
(SOLPT) populations in response to sulfosulfuron treatment .................................. 62

Table 10. Log logistic equations, R2 values and GRSO values for eastern black nightshade
(SOLPT) populations in response to pyridate treatment63

Table 11. Log logistic equations, R2 values and GR50 values for hairy nightshade
(SOLSA) populations in response to metribuzin treatment .................................... 64

Table 12. Log logistic equations, R2 values and GRso values for hairy nightshade
(SOLSA) populations in response to sulfentrazone treatment .................................. 65

Table 13. Log logistic equations, R2 values and GR50 values for hairy nightshade
(SOLSA) populations in response to sulfosulfuron treatment .................................. 66

Table 14. Log logistic equations, R2 values and GR50 values for hairy nightshade
(SOLSA) populations in response to halosulfuron treatment .................................. 67

Table 15. Log logistic equations, R2 values and GRSO values for hairy nightshade
(SOLSA) populations in response to pyridate treatment.............................. ...........68

Table 16. Log logistic equations, R2 values and GR50 values for horsenettle (SOLCA)
populations in response to halosulfuron treatment .............................................. 69

Table 17. Log logistic equations, R2 values and GR50 values for horsenettle (SOLC A)
populations in response to sulfentrazone treatment .............................................. 70

viii

Table 18. Log logistic equations, R2 values and GR50 values for horsenettle (SOLCA)
populations in response to sulfosulfuron treatment .............................................. 71

Table 19. Log logistic equations, R2 values and GRSO values for horsenettle (SOLCA)
populations in response to pyridate treatment .................................................... 72

Table 20. Log logistic equations, R2 values and GR50 values for horsenettle (SOLCA)
populations in response to metribuzin treatment ................................................ 73

Table 21. Log logistic equations, R2 valuesa and GR50 values for clammy groundcherry
(PHYHE) population in response to sulfosulfuron, metribuzin, pyridate, sulfentrazone
treatment ............................................................................................... 74

Table 22. Log logistic equations, R2 values3 and GRso values for smooth groundcherry
(PHYSU) population in response to sulfosulfuron, pyridate, sulfentrazone treatment... .75

Table 23. Range of GR50 values between populations of eastern black nightshade

(SOLPT), hairy nightshade (SOLSA), horsenettle (SOLCA), clammy groundcherry
(PHYHE) and smooth groundcherry (PHYSU) in response to different treatments ....... 76

CHAPTER 4

EFFECT OF TEMPERATURE ON GERMINATION OF EASTERN BLACK
NIGHTSHADE, HAIRY NIGHTSHADE, HORSENETTLE, SMOOTH
GROUNDCHERRY AND CLAMMY GROUNDCHERRY

Table 1. Comparison of germination mean percent across temperature to analyze the
trend in variability among eastern black nightshade (SOLPT) populations, horsenettle
(SOLCA) populations, hairy nightshade (SOLSA) population, smooth groundcherry
(PHYSU) and clammy groundcherry (PHYHE) species ...................................... 122

Table 2. List of eastern black nightshade, hairy nightshade, horsenettle, clammy
groundcherry and smooth groundcherry populations collected from various regions in
Michigan, its location, associated crop and seed viability percent .......................... 123

LIST OF FIGURES

CHAPTER 2
POTENTIAL HERBICIDES TO CONTROL EASTERN BLACK NIGHTSHADE
(Solan um ptycanth um) IN TOMATO (Lycopersicon esculentum)

Figure 1. Polynomial quadratic comparison Of eastern black nightshade (SOLPT)
response to sulfosulfuron at 0.0086 kg/ha, 0.0173 kg/ha, 0.034 kg/ha, 0.069 kg/ha and
0.13 kg/ha concentrations applied during early 2003 Spring and mid summer season in
greenhouse......... ............................................................................................................. 38

Figure 2. Polynomial quadratic comparison of eastern black nightshade (SOLPT)
response to sulfentrazone at 0.028 kg /ha, 0.056 kg /ha, 0.11 kg ai/ha, 0.22 kg /ha, and
0.44 kg /ha concentrations applied during early 2003 spring and mid summer season in
greenhouse.......... ............................................................................................................ 39

CHAPTER 3
VARIATION IN HERBICIDE RESPONSE AMONG NIGHTSHADE (Solanum
spp) AND GROUNDCHERRY (Physalis spp) SPECIES AND WITHIN
NIGHTSHADE POPULATIONS

Figure 1a. Log logistic dose response curves Of eastern black nightshade populations -
Monroe4, Masonl , Monroe] , and NewagOZ in response to metribuzin ..................... 77

Figure 1b. Log logistic dose response curves of eastern black nightshade populations -
Macombl , Macomb2, Oceanal, and Macomb3 in response to metribuzin .................. 78

Figure 1c. Log logistic dose response curves of the most susceptible - Monroe4 and
tolerant - Oceanal eastern black nightshade populations in response to metribuzin ...... 79

Figure 2a. Log logistic dose response curves of eastern black nightshade populations -
NewagOZ, Macomb3 , Macomb 1 , and Macomb2 in response to
sulfentrazone.................. .............................................................................................. 80

Figure 2b. Log logistic dose response curves of eastern black nightshade populations -
Oceanal , Monroe4, Monroe2, and Masonl in response to sulfentrazone ................... 81

Figure 2c. Log logistic dose response curves Of eastern black nightshade populations -
Bayl , Monroel, Ingham2, and Inghaml in response to sulfentrazone ..................... .82

Figure 2d. Log logistic dose response curves of the most susceptible NewagoZ and
tolerant Ingham2 eastern black nightshade populations in response to sulfentrazone. . .. .83

Figure 3a. Log logistic dose response curves of eastern black nightshade populations -
Macomb3, Macombl, Monroe4, Oceana], and Monrer in response to
halosulfuron......... ........................................................................................................... 84

Figure 3b. Log logistic dose response curves of eastern black nightshade populations —
Monroel, Inghaml , Newag02, Monroe2, and Bay] in response to halosulfuron ......... .85

Figure 3c. Log logistic dose response curves of the most susceptible Macomb3,
Macombl and tolerant Newag02, Macomb2 eastern black nightshade populations in
response to halosulfuron ............................................................................ 86

Figure 4a. Log logistic dose response curves of eastern black nightshade populations -
Monroe2, Monroe], Bay] , and Macomb] in response to sulfosulfuron ..................... 87

Figure 4b. Log logistic dose response curves of eastern black nightshade populations —
Macombl, Monroe4, Inghaml , and Newag02 in response to sulfosulfuron ............... 88

Figure 4c. Log logistic dose response curves of eastern black nightshade populations -
Mason] , Ingham2, Macomb3, and Oceanal in response to
sulfosulfuron.............................. 89

Figure 4d. Log logistic dose response curves of the most susceptible Macomb2 and
tolerant Oceanal eastern black nightshade populations in response to sulfosulfuron. .. ...90

Figure 5a. Log logistic dose response curves of eastern black nightshade populations -
Masonl, Ingham2, Inghaml, Bay] and Oceanal in response to
pyridate........................ .............................................................................................. 91

Figure 5b. Log logistic dose response curves of hairy nightshade populations -
Montcalm], Montcalm2, Presque2, and Presque3 in response to metribuzin ............... 92

Figure5b. Log logistic dose response curves of hairy nightshade populations - Presque4,
Presquel , Macombl, and Bay2 in response to metribuzin .................................... 93

Figure Sc. Log logistic dose response curves Of the most susceptible Montcalmland
tolerant Macomb] hairy nightshade populations in response to metribuzin. . . . . . . . . . .. .....94

Figure 6a. Log logistic dose response curves of hairy nightshade populations -Presque2,
Presque3 and Presquel, Montcalm2 in response to sulfentrazone. . . . . ...............95

Figure 6b. Log logistic dose response curves of hairy nightshade populations - Presque4,
Montcalm] , Macombl , and Bay2 in response to sulfentrazone .............................. 96

Figure 6c. Log logistic dose response curves of the most susceptible Presque2 and
tolerant Macombl hairy nightshade populations in response to sulfentrazone. . . . . . . .. .....97

xi

Figure 7a. Log logistic dose response curves of hairy nightshade populations - Presque2,
Presque3, and Presque4 in response to sulfosulfuron .......................................... 98

Figure 7b. Log logistic dose response curves of hairy nightshade populations - Macombl ,
Montcalm] , and Bay2 in response to sulfosulfuron ............................................ 99

Figure 70. Log logistic dose response curves Of the most susceptible Presque4 and
tolerant Macomb] hairy nightshade populations in response to sulfosulfuron ............ 100

Figure 8a. Log logistic dose response curves Of hairy nightshade populations - Presque2,
Presque3, Montcalm2, Presque4, Bay2 and Macomb] in response to halosulfuron ...... 101

Figure 8b. Log logistic dose response curves of the most susceptible Presque2 and

tolerant Macombl hairy nightshade populations in response to halosulfuron ............ 102
Figure 9. Log logistic dose response curves of horsenettle populations - Berrien2,
Vanburenl, Montca1m3, and Monroe5 in response to halosulfuron ........................ 103
Figure 10. Log logistic dose response curves of horsenettle populations - Berrien2,
Vanburenl, Montca1m3, and Monroe5 in response to sulfentrazone ....................... 104
Figure 11. Log logistic dose response curves of horsenettle populations - Berrien2,
Montca1m3, and Monroe5 in response to sulfosulfuron........................... ..............105
Figure12. Log logistic dose response curves of horsenettle populations - Berrien2,
Vanburenl, Montca1m3, and Monroe5 in response to pyridate .............................. 106

CHAPTER 4

EFFECT OF TEMPERATURE ON GERMINATION OF EASTERN BLACK
NIGHTSHADE, HAIRY NIGHTSHADE, HORSENETTLE, SMOOTH
GROUNDCHERRY AND CLAMMY GROUNDCHERRY

Figure 1. Temperature effect on the period of germination of Oceanal eastern black
nightshade population at A) 28/200 C, B) 25/160 C, C) 22/140 C and D) 15/ 100 C ...... 124

Figure 2. Temperature effect on the period Of germination Of Inghaml eastern black
nightshade population at A) 28/20 0C, B) 25/ 16 °C and C) 22/14 °C ..................... 125

Figure 3. Temperature effect on the period of germination of clammy groundcherry
population Newagol at A) 24/16 0C, B) 28/20 OC and smooth groundcherry population
Ingham3 at C) 22/14 °C, D) 24/16 0C, E) 28/20O C .......................................... .126

Figure 4. Temperature effect on the period Of germination of Vanburen] horsenettle
population at A) 28/20 °C, B) 25/16 0C, and C) 22/14 0C ................................... 127

xii

Figure 5. Temperature effect on the period Of germination Of Oceana4 horsenettle
population at A) 28/20 °C, B) 25/16 0C, and C) 22/14 0C ............................................... 128

xiii

Figure 5. Temperature effect on the period of germination of Oceana4 horsenettle
population at A) 28/20 0C, B) 25/16 °C, and C) 22/ 14 0C ............................................... 128

 

xiii

CHAPTER 1

LITERATURE REVIEW

Introduction:

Eastern black nightshade (Solanum plycanthum Dun.) is native to North America
and is found mostly east of the Rocky Mountains (Ogg et al., 1981). Eastern black
nightshade is one of the most troublesome weeds in soybean (Glycine max 1.), tomato
(Lycopersicon esculentum L.), pepper (Capsicum annuum 1.), ﬁeld bean (Phaseolus
vulgaris L.), and corn (Zea mays L.) in central and north eastern America (Ogg et al.,
1981). In Nebraska, economic losses due to eastern black nightshade infestation in
soybean ﬁelds have been estimated at 12% of total state soybean income (Burgert et al.,
1973). Eastern black nightshade causes a major problem in soybean harvesting operations
by forming a sticky mass which can quickly plug combine screens and rotors. Mature
berries Often stain soybean during harvest and thereby reduce the bean quality. Eastern
black nightshade causes a major problem in fresh pea harvesting operations. Immature
berries of eastern black nightshade are similar in size, shape and color to pea, which
makes separation of pea and nightshade fruit difﬁcult during harvest. During processing,
the immature berries turn black and reduce the pea quality (Majek, 1981). Perez and
Masiunas (1990) estimated 60% yield loss in transplanted tomato with four eastern black
nightshade plants per m2. Also, eastern black nightshade is an alternate host for many

tomato pests and diseases.

Taxonomic characteristics:

The Solanum nigrum (nightshade) complex consists of approximately 30 species.
In the US there are I] recognized species (Schilling, 1981) of Solanum with four species
in the Solanum nigrum complex recognized as troublesome weeds in a variety Of crops
(Heiser et a], 1979). These species include hairy nightshade (Solanum sarrachoides
Sendtner), eastern black nightshade, black nightshade (Solanum nigrum) and American
black nightshade (So/anum americanum Mill.) (Ogg et al., 1981). These closely related
species are difﬁcult to separate in taxonomic classiﬁcation. There are four primary
reasons for difﬁculty in taxonomic identiﬁcation and separation of the species (Ogg et al.,
1981). First and most Obvious, is the similarity in gross morphology among species.
Second, these species are highly phenotypically plastic. When grown under different
environmental conditions, they vary considerably in many taxonomic characteristics that
are frequently used for identiﬁcation. Third, genetic variation in certain species is large
and is expressed as numerous geographic types within a single species. Fourth, there has
been extensive nomenclature confusion within the group. Hence there is a need to
understand some of the key taxonomic characteristics of eastern black nightshade.

The following description of eastern black nightshade was based on work by
Bassett & Munro (1984), Ogg et al., (1981), and Schilling (1981).
Eastern black nightshade (Chromosome no. 2n= 24):

Eastern black nightshade is an annual, rarely a short-lived perennial, up to 1m
high, erect, sparsely or freely branching plant. The stems are green or greenish purple,
round or angular, and subglabrous. The lower surfaces of the young seedling leaves are

purple in color. Leaves are up tO 10 cm long and 9 cm broad, triangular-ovate or elliptic

and acute. The surfaces are subglabrous with few eglandular hairs. The inﬂorescence is
umbellate, up to six ﬂowers and the calyx is 1-3 mm long at anthesis and the lobes adhere
to the mature berry. Corolla is stellate, white with a yellow star, about 8 mm broad.
Anthers are 1-2 mm long and yellow. Mature berries are globosely, shiny, and purplish
black. The berries contain 6-15 sclerotic granules and up to 100 seeds. The seeds are 1.5 -
1.8 mm long and 1.3 mm wide, light brown in color.

Eastern black nightshade is most frequently confused with American black
nightshade and black nightshade because of their similar appearance.
History:

Eastern black nightshade is a native Of North America and was falsely identiﬁed
in Michigan as black nightshade until the 1980’s (Vandeventer et al., 1982). Eastern
black nightshade was ﬁrst collected in Michigan in 1896 in Grand Rapids by J.S.
Haddick and H.C.Skeels (MSU herbarium).

Seed production and dispersal characteristics:

The time period from germination to fruiting of eastern black nightshade can be
completed within 6 weeks; however, ﬂowering can last for several months (Ogg et al.,
1981). Eastern black nightshade produces viable seeds within 4 weeks of anthesis
(Quakenbush and Anderson, 1984). Eastern black nightshade plants can produce many
berries and viable seeds within a short period of time. Each eastern black nightshade
plant produces 50 to 100 berries (Bassett and Munro, 1984) and each berry may contain
110 to 150 seeds (Majek, 198]; Roberts and Lockett, 1978). In another study, Stoller and
Myers (1989a) reported that eastern black nightshade produced 50,000 seeds per plant

under full sunlight and 20,000 seeds per plant with soybean interference. Eastern black

nightshade seeds have viability Of 90% in soil for 5 years. Viability drops successively to
73, 27 and 2% over the next three years (Dorph-Petersen, 1924).

Berries and seeds of eastern black nightshade are easily dispersed by rodents,
birds, livestock, man and along water courses (Salisbury, 1961; Burgert et al., 1973;
Kelley & Bruns, 1975; Roberts & Lockett, 1978). During soybean harvesting operations,
berries are crushed and the sticky seeds are easily disseminated by the harvesting
equipment throughout the ﬁeld.

GERMINATION CHARACTERISTICS:

The invasion success of a colonizing weed species depends on the ability of its
seed to germinate in new habitats (Groves, 1986). Many germination studies have been
done to estimate the colonization potential of eastern black nightshade and to predict time
of emergence in the ﬁeld. Hermanutz and Weaver (1990) Observed that the ruderal
eastern black nightshade populations of northern range germinated faster than agrestal
eastern black nightshade populations, though both populations had similar base
temperatures. This genetic variability in germination parameters across temperature may
have profound impact on future range of expansion. Hermanutz and Weaver (1991)
Observed that eastern black nightshade germinates more slowly over a wider range of
temperature than hairy nightshade, and may have prolonged germination during the
growing season. Hence, eastern black nightshade can produce more seeds and seedlings
per year and can be more dominant than hairy nightshade with respect to colonization.

Reports in general suggest that alternating temperature from 20 to 30 C causes
maximum germination of eastern black nightshade. However under darkness, very poor

germination Of eastern black nightshade occurs (Thomson and Witt, 1987; Hermanutz

and Weaver, 1990). Germination of eastern black nightshade is region speciﬁc and varies
under different environmental conditions (Hermanutz and Weaver, 1990). In general,
eastern black nightshade emerges in the late spring and continues until mid summer when
the daily maximum air temperature approaches 20 C (Ogg and Dawson, 1984; Roberts
and Lockett, 1978). In addition, shallow tillage can stimulate overall emergence of
eastern black nightshade in ﬁeld conditions (Ogg and Dawson, 1984).

Growth and physiological characteristics:

Eastern black nightshade has a dichotomous branching pattern, but the forks are
unequal in length. This unequal fork results in a spreading growth habit for eastern black
nightshade and greater leaf area (McGiffen et al., 1992) and thereby it forms a dense
canopy to intercept more light (McGiffen & Masiunas, 1992). The plant also responds to
self-shading by increasing speciﬁc leaf area, and forms thinner leaves. Thinner leaves can
result in decreased respiration rate and more carbon assimilation per quantum of light
(Stoller & Myers, 1989b). Thus, by adapting to self-shading, eastern black nightshade
maintains a dense canopy that can shade the growth of other plants and increase the
efﬁciency Of competitiveness.

In addition to self-shading, eastern black nightshade also responds to the shading
effect Of the crop canopy by allocating more biomass to leaves and less biomass to
berries, lowering respiration rate, and increasing speciﬁc leaf area and light absorption
efﬁciency. This physiological adaptation to shade increases the efﬁcient utilization of

photosynthetic active radiation (PAR) (Stoller & Meyer, 1989a).

Interference in tomato growth and development:

Tomato is one of the most important vegetable crops in Michigan. Bridges (1992)
estimated the annual tomato yield loss due to weeds in Michigan was about 6.4 million
dollars in fresh market tomatoes and 11.9 million dollars in processing tomatoes. Eastern
black nightshade is one of the major competitive weeds in Michigan tomatoes.

Reports Of tomato yield loss from nightshade interference vary widely. In
southern France, 20 black nightshade plants per meter of row caused 73% reduction in
transplanted tomato yield (Maillet and Abdel-Fatah, 1983). In southern Italy, eight black
nightshades plants per meter caused similar yield reduction of about 73% (Damato and
Montemurro, 1986). And in Canada, a mixture of 20 eastern black nightshades and hairy
nightshade plants per meter of row decreased transplanted tomato yield by 20 to 60% and
direct-seeded tomato yield by about 95% (Weaver et al., 1987).

Perez and Masiunas (1990) observed that the critical stage for control of eastern
black nightshade is within 6 weeks after tomato transplantation. Eastern black nightshade
begins to interfere with tomato from 4 to 8 weeks after tomato transplanting (Perez &
Masiunas, 1990; Weaver and Tan, 1983). This critical period coincides with the full
bloom stage of tomato (F riesen, 1979; Perez & Masiunas, 1990). So, if unchecked within
6 weeks, three eastern black nightshade plants per m2 can reduce tomato yield by 60%.
During this critical period of tomato interference, eastern black nightshade grew much
faster and taller than tomato, competing for light by overtopping tomato’s canopy
(Weaver and Tan, 1983). Thus eastern black nightshade reduces the availability of PAR

to tomato, which is necessary for its ﬂower and fruit development.

Eastern black nightshade also competes with tomato for water and nutrients.
Wahle and Masiunas (2003), Observed that eastern black nightshade responds to high
levels Of nitrogen (N). Eastern black nightshade attained maximum growth at 12 weeks
after planting and fresh mass increased with N up to 336 kg N/ha. Similar Observations
were reported by Bassett and Munro (1985), and Gonzalec et al., (1996). Thus eastern
black nightshade can be more competitive with tomato in accumulation Of N for its
growth and development.

Tan and Weaver (1996) reported that eastern black nightshade possesses more
stomata (particularly on the lower leaf surface) than hairy nightshade. This causes an
increase in transpiration rate under ample water and light conditions. Hence, due to high
transpiration rate, eastern black nightshade competes strongly with tomato for water
which may cause rapid depletion Of soil water resources.

Alternate host to pest and diseases:

Nightshade acts as an alternate host to many pests and diseases in Solanaceous
crops. Colorado potato beetle (Leptinotarsa decemlineata Say) feeds on nightshades
(Brown et al., 1980). Rhizoctonia solani frequently colonizes the roots of nightshade in
potato ﬁelds, contributing to the transmission of the disease from one season to another
(Ogg, 1989). Nightshades are an excellent host of late blight (Phytophthora infestans)
(Vartanian and Ends, 1985) and powdery mildew (Oidium lycopersicum) (Lamondia et
al., 1999) which are common diseases in tomato. Perennial eastern black nightshade acts
as an overwintering host for cucumber mosaic virus in pepper (Hobbs et al., 2000).
Eastern black nightshade can also host nematodes like Heterodera glycines (Wong and

Tylka., 1994), and Globodera tabacum (Lamondia, 1996).

Difficult to control in tomato:

Eastern black nightshade and tomato have similar herbicide susceptibilities,
growth habitat and physiology (McGiffen and Masiunas, 1991; Perez and Masiunas,
1990; Weaver et al., 1987). Hence it is difﬁcult to ﬁnd a selective herbicide to control
eastern black nightshade without injuring tomato. Currently, there are few registered
herbicides for use in tomato which limits the control options of weeds like nightshades.
Metolachlor is currently registered for use in tomato and gives good control of eastern
black nightshade preemergence (Gaynor et al., 1993). Currently there are no registered
herbicides available which will control eastern black nightshade postemergence in
tomato.

Metribuzin is registered for broadleaf weed control in tomato, but does not control
eastern black nightshade (Ackley et al., 1997). A combination of metribuzin and pyridate
controlled eastern black nightshade, but caused unacceptable injury to tomato (McGiffen
and Masiunas, 1991). DCPA and chloramben herbicides are no longer registered for use
in tomato due to a potential leaching effect in the soil (Perez and Masiunas, 1990).
Rimsulfuron, a sulfonylurea herbicide, is used in tomato production worldwide (Reinke
et al., 1991), but lacks persistence and does not control black nightshades. Some tomato
cultivars are also sensitive to rimsulfuron (Bewick et al., 1991).

Chang and Masiunas (1992) characterized some of the somoclones of eastern
black nightshade tolerant to aciﬂuorfen, oxyﬂuorfen, diquat and paraquat and suggested
the tolerance of nightshade to the diphenyl ether and bipyridylium salt herbicides could
be a problem in ﬁeld situations. The reliance on a few active ingredients (and sites Of

action) for nightshade management in ﬁeld crops has resulted in herbicide resistance

problems. Biotypes of eastern black nightshade resistant to acetolactate synthesis (ALS)
inhibitor herbicides such as imazethapyr, imazamox and primisulfuron-methyl have been
reported in Wisconsin (Volenberg et al., 2000), Indiana, and Illinois (Milliman et al.,
2003)

Many fresh market tomato growers have adopted the plasticulture production
system to achieve earlier harvest and increase high quality fruit yield (Brown et al., 1991;
Wein and Minotti, 1987) and to reduce the weeds. In a plasticulture production system,
fumigation by methyl bromide has been successfully used to control many Weeds. The
phaseout of methyl bromide by 2005 in the US (EPA, 1998), has further limited the
Options for weed control in tomatoes. Hence, there are no alternate methods to control

eastern black nightshade in tomato.

LITERATURE CITED

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efﬁcacy in transplanted tomato (Lycopersicon esculentum). Weed Technology
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ptycanthum Dun, S. nigrum L. and S.sarrach0ides Sendt. Canadian Journal Of
Plant Science 65:401-414.

. Bewick, T.A., W.M. Stall, S.R. Kostewicz, and K. Smith. 199]. Alternatives for
control of paraquat tolerant American black nightshade (Solanum Americana).
Weed Technology 5:61-65.

. Bridges.1992. Crop losses due to weeds in the United States. Allen Press,
Lawrence, KS pg30.

. Brown, J.E., W.D. Goff, W. Hogue, M.S. West, C. Stevens, V.A. Khan, B.C.
Early and LS. Brasher. 1991. Effects of plastic mulch on yield and earliness of
tomato. Proceeding of Natural Agriculture Plastics Conference 23:21-28.

. Brown, J ., T. Jeremy and BA. Butt. 1980. The inﬂuence of an alternative host
plant on the fecundity Of the Colorado potato beetle (Leptinotarsa decemlineata).
Annual Entomology Society Of American 73:197-199.

. Burgert, K.L., Brunside, CC. and Fenster, CR. 1973. Black nightshade (Solanum
nigrum) leaves its mark. Nebraska University College of Agriculture. Serving
farm 20:8-10.

. Chang Y.Y. and J .B. Masiunas, 1992. Characterization of aciﬂuorfen tolerance in
selected somaclones of eastern black nightshade (Solanum ptycanthum). Weed

Science 40:408-412.

. Damato, G. and P. Montemurro. 1986. Studio della competizione fra Solanum
nigrum L. e pomodoro da industria trapianto. La Difesa Delle Piante 9:359-364.

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station during the years 1896-1923. International Seed Test Congress Rep.4:124-
138.

EPA (U .8. Environmental Protection Agency).1998. Methyl bromide information
In: Ozone depletion. (1998, NOV. 10), [Online] Available:
http://www.epa.gov/ozone/mbr/mbrqa.html., Dec.] 1998.

Friesen, G.H. 1979. Weed interference in transplanted tomatoes (Lycopersicon
esculentum). Weed Science 27: 1 1-13.

Gaynor, J.D., Hamil, A.S., MacTavish, DC. 1993. Efﬁcacy, fruit residues and
soil dissipation Of the herbicide metolachlor in processing tomato. Journal of
American Society of Horticulture Science 118:68-72.

Gonzalec P., Zancada C. Vedugo M., and Salas L. 1996. Plant height as a factor
in competition between black nightshade and two horticultural crops (tomato and
pepper). Journal Of Horticulture Science 71:.453-460

Groves, RH. 1986. Invasion of Mediterranean ecosystems by weeds. In:
Resilience in Mediterranean- type ecosystems. Edited by B. Dell, A.J.M. Hopkins
and BB. Lamont. Junk Publishers, Boston, MA pp.129-145.

Heiser, C.B., Burton, BL. and Schilling, E.E, Jr. 1979. Biosystematics and
taxonomic studies of the Solanum nigrum complex in eastern North America.
The biology and taxonomy Of the Solanaceae. Academic Press, NY pg 513-527.

Hermanutz, LA. and SE. Weaver. 1990. Variability in temperature- dependent
germination in eastern black nightshade (Solanum ptycanthum). Canadian J. Bot.
69:463-1470.

Hermanutz, LA. and SE. Weaver. 1991. Germination and growth of Solanum
plycanthum and Solanum sarrachoides. Canadian Journal Of Plant Science 71:39-
40].

Hobbs, H.A., D.M. Eastbum, C.J.D. Arcy, J.D. Kindhart. J.B. Masiunas, D.J.
Voegtlin, R.A. Weinzierl and N.K. McCOppin. 2000. Solanaceous weeds as

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possible source Of Cucumber mosaic virus in southern Illinois for aphid
transmission to pepper. Plant Disease 84: 1221-1224.

Kelley, AD. and Bruns, V.F. 1975. Dissemination Of weed seeds by irrigation
water. Weed Science 23:486-493.

Larnondia, J.A. 1996. Trap crops and population management Of Globodera
tabacum tabacum. Journal Of Nematology 28:238-243.

Lamondia, J.A., Smith, V.L., Douglass, SM. 1999. Host range Of Oidium
chopersicum on selected Solanaceous species in Connecticut. Plant Disease
83:341-344.

Maillet J. and H.Abdel-Fatah. 1983. Etudes preliminaries sur la concurrence entre
Solanum nigrum spp. Eu-nigrum L. (morelle moire) et Lycopersicon esculentum
Mill. (tomate) enculture repiquee. Weed Research 23:217-219.

Majek, BA. 1981. Nightshade control in ﬁeld crops. University Of Minnesota
Agricultural Extension folder 603:2-4.

McGiffen, Jr., ME. and J.B. Masiunas. 1991. Post-emergence control Of
broadleaf weeds in tomato. Weed Technology 5:739-745.

McGiffen, Jr., ME. and J.B. Masiunas. 1992. Prediction of black and eastern
black nightshade (Solanum nigrum & S. ptycanthum) growth using degree-days.
Weed Science 40:220-226.

McGiffen, Jr., M.E., J.B. Masiunas and J. Hesketh. 1992. Competition for light
between tomatoes and nightshades (Solanum nigrum and S.ptycanthum). Weed
Science 40:220-226.

Milliman, L.D., D.E. Riechers, L.M. Wax and F.W. Simmons. 2003.
Characterization of two biotypes of imidazolinone — resistant eastern black

nightshade (Solanum ptycanthum). Weed Science 51:139-144.

12

29. Ogg A.G., Jr., B.S. Rogers and BE. Schilling. 1981. Characterization of black

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nightshade (Solanum nigrum) and related species in the United States. Weed
Science 29:27-32.

Ogg, A.G., Jr. and J .H. Dawson. 1984. Time of emergence of eight weed species.
Weed Science 32:327-335.

Ogg, A.G., Jr. 1989. Taxonomy, distribution, biology and control of black
nightshade (Solanum nigrum) and related species in United States and Canada.
Rev. Weed Science 4:25-58.

Perez, F .G.M. and J.B. Masiunas. 1990. Eastern black nightshade (Solanum
ptycanthum) interference in processing tomato (Lycopersicon esculentum). Weed
Science 38:385-388.

Quackenbush LS. and RN. Andersen. 1984. Distribution and biology of two
nightshades (Solanum spp.) in Minnesota. Weed Science 32:529-533.

Reinke, H., Rosenzweig A. K.M. Clausm, C.Chisholm and P.Jensen. 1991. DPX-
E9636, experimental sulfonylurea herbicide for potatoes. Proceeding Of Brighton
Crop Protection Conference-Weeds 445-451.

Roberts, HA. and Lockett, P.M.1978. Seed dormancy and ﬁeld emergence in
Solanum nigrum. Weed Research 18:231-241.

Salisbury, E. 1961. Weeds and aliens. Collins, London pp384.

Schilling, BE. 1981. Systematics of Solanum sect. Solanum (Solanaceae) in
North America Systematic Botany 6:172-185.

Stoller, E.W.and R.A. Myers. 1989a. Effects on shading and soybean (Glycine
max L.) interference in eastern black nightshade (Solanum ptycanthum) growth
and development. Weed Research 29:307-317.

39. Stoller, E.W. and RA. Myers. 1989b. Response of soybeans (Glycine max L.) and

four broadleaf weeds to reduced irradiance. Weed Science 37:570-574.

13

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Tan CS. and SE. Weaver, 1996. Water use patterns of eastern black nightshade

and hairy nightshade in response to shading and water stress. Canadian Journal
Plant Science 77:261-265.

Thomson, EC. and W. Witt. 1987. Germination Of cutleaf groundcherry (Physalis
angulata), smooth groundcherry (Physalis virginiana) and eastern black
nightshade (Solanum ptycanthum). Weed Science 35:58-62.

Vandeventer, J.W., W.F. Meggitt, and D.Penner.l982. Morphological and

physiological variability in black nightshade (Solanum spp) Pesticide Science
13:257- 262.

Vartanian, V.G. and RM. Ends.1985. Overwintering hosts, compatibility types

and races of Phytophthora infastans on tomato in Southern California. Plant
disease 69:516-519.

Volenberg S., Stoltenberg E., Boerboom M. 2000. Solanum ptycanthum resistance
to acetolactate synthase inhibitors. Weed Science 482399-401

Wahle, EA. and J.B. Masiunas. 2003. Comparison of nitrogen use by two
population densities of eastern black nightshade (Solanum ptycanthum). Weed
Science 51:394-401.

Weaver, SE. and CS. Tan. 1983. Critical period Of weed interference in
transplanted tomatoes (Lycopersicon esculentum): growth analysis. Weed Science
31:467-481.

Weaver, S.E., N. Smits, and CS. Tan. 1987. Estimating yield losses of tomatoes
(Lycopersicon esculentum) caused by nightshade (Solanum spp.) interference.
Weed Science 35:163-168.

Wein, HQ and PL. Minotti. 1987. Growth, yield, and uptake Of transplanted
fresh-market tomato as affected by plastic mulch and initial nitrogen rate. Journal
Of American Society Horticulture Science 112:759-763.

Wong-ATS, Tylka G.L. 1994. Eight non-host weed species of Heterodera
glycines in Iowa. Plant Disease 78:365-367.

14

CHAPTER 2
POTENTIAL HERBICIDES TO CONTROL EASTERN BLACK NIGHTSHADE
(Solanum ptycanth um) IN TOMATO (L ycopersicon esculentum)
Abstract. Eastern black nightshade (Solanum ptycanthum Dun.) is a troublesome weed in
tomato production. Eastern black nightshade and tomato share similar herbicide
susceptibilities due to a close genetic relationship. The Objective of this study was to
evaluate potential herbicides to control eastern black nightshade in tomato. Field studies
showed that post-transplant application Of S-metalochlor (1.8 kg/ha), dimethenamid-P
(1.1 kg/ha) and pre-transplant application Of ﬂumioxazin (0.052 kg/ha), and oxyﬂuorfen
(0.28 kg/ha) gave at least 95% control Of eastern black nightshade with negligible or no
crop injury and no yield reduction. Postemergence application of pyridate controlled 60%
control of eastern black nightshade with no crop injury and resulted in yield of 85% of
the weeded control. Pre-transplant and postemergence application of sulfentrazone gave
fair control of eastern black nightshade, but caused crop injury of at least 20%. Post-
directed application of carfentrazone (0.18 kg/ha) and ﬂumioxazin (0.052 kg/ha) caused
unacceptable crop injury to tomato. In greenhouse studies, sulfosulfuron (0.034 kg/ha)
did not control eastern black nightshade sufﬁciently and did not cause injury to tomato.
However, eastern black nightshade has shown a difference in response to sulfosulfuron
applied during the spring and summer seasons. Eastern black nightshade showed more
sensitivity to sulfosulfuron during the spring season than during the summer season.
Key words: Tomato, eastern black nightshade (Solanum ptycanthum), dimethenamid-P,
S-metolachlor, sulfosulfuron, ﬂumioxazin, oxyﬂuorfen, sulfentrazone, pyridate,

carfentrazone.

15

INTRODUCTION

Tomato (Lycopersicon esculentum Mill.) is one of the most important vegetable crops in
Michigan. Interference Of weeds in tomato causes yield reduction and low quality of
tomatoes (Friesen, 1979; McGiffen et al., 1992). Eastern black nightshade (Solanum
ptycanthum Dun.) is a troublesome weed in tomato throughout the northeastern United
States. Eastern black nightshade interferes with tomato within 4 to 8 weeks after crop
establishment (Perez and Masiunas, 1990; Weaver and Tan, 1983), coinciding with the
full bloom stage of tomato (F riesen, 1979). Eastern black nightshade overtops the tomato
canopy resulting in a decrease photosynthetic active radiation (PAR), ﬂower abortion and
less fruit set (Mc Giffen et al., 1992). Four eastern black nightshade plants per meter
square can cause a yield reduction of 80% in direct seeded tomatoes and 25 to 60% in
transplanted tomatoes (Perez and Masiunas, 1990).

Eastern black nightshade is difﬁcult to control in tomato because eastern black
nightshade and tomato share a close genetic relationship, having similar herbicide
susceptibilities, growth habitat and physiology (McGiffen et al., 1992; Perez and
Masiunas, 1990; Weaver et al., 1987). There are few registered herbicides in tomato
(Ackley et al., 1997). Metolachlor is currently the only registered herbicide to control
eastern black nightshade preemergence in tomato and no postemergence herbicides are
registered for eastern black nightshade control in tomato.

Metribuzin is registered for control of broadleaf weeds in tomato but it doesn’t
control eastern black nightshade (Ackley et al., 1997). Combinations of metribuzin and
pyridate controlled eastern black nightshade, but caused unacceptable injury to tomato

(McGiffen and Masiunas, 1991). DCPA and chloramben are no longer registered for use

16

in tomatoes (Perez and Masiunas, 1990). Rimsulfuron, a sulfonylurea herbicide is used in
tomato production worldwide (Reinke et al., 1991), but it lacks persistence and does not
control black nightshade. Besides, some tomato cultivars are sensitive to rimsulfuron
(Bewick et al., 1995).

The reliance on a few active ingredients (and sites of action) for nightshade
management in ﬁeld crops has resulted in herbicide-resistant weed problems. Biotypes of
eastern black nightshade resistant to ALS inhibitor herbicides such as imazethapyr,
imazamox and primisulfuron-methyl have been reported from Wisconsin (V Olenberg et
al., 2000), Indiana and Illinois (Milliman et al., 2003).

Many fresh market tomato growers have adopted the plasticulture production
system to achieve earlier harvest, increase high quality fruit yield (Brown et al., 1991;
Wein and Minotti, 1987) and to reduce weeds. In the plasticulture production system,
fumigation by methyl bromide successfully controls many weeds. The phase out of
methyl bromide by 2005 in the US (EPA, 1998) has further limited the Options for
control of eastern black nightshade in tomatoes.

Due to the prevalence and strong interference Of eastern black nightshade in
tomato growth and development, and the necessity for alternate herbicides to establish an
effective management system in tomato, ﬁeld studies were conducted to evaluate the

efﬁcacy of potential herbicides to control eastern black nightshade in tomato.

17

MATERIALS AND METHODS
FIELD STUDY:

Field experiments were conducted in 2002, 2003 and 2004 to evaluate the
potential herbicides to control eastern black nightshade in tomato. The research was
conducted at Michigan State University’s Horticulture Teaching and Research Center
(HTRC), East Lansing, Michigan. The soil type at the research center was Marlette ﬁne
sandy loam. In 2002 and 2004, the experimental site had very high eastern black
nightshade pressure. The 2003 site had low to moderate eastern black nightshade pressure
but there was sufﬁcient pressure for herbicide evaluation.

Field preparation:

The ﬁelds were plowed using a moldboard plow at the end of the previous fall
season. In mid spring, the ﬁelds were worked with a ﬁeld cultivator to enhance the soil
aeration and decomposition of the previous residues. The ﬁeld was fertilized with N:
P205:K20 ratio of 19: 19:19 at rates Of 250 kg/ha, 312 kg/ha, 300 kg/ha in 2002, 2003 and
2004 respectively.

Transplanting:

The tomato cultivar Pikritel was used in this research. The seeds were sown in
ﬂats in the greenhouse and were grown for about 30 days and then transferred to a shade
house for 10 days before being transplanted into the ﬁeld. Forty-day-old tomato seedlings
were transplanted with a Mechanical Transplanter Model 40002 with two rows 90cm
apart per plot with 60 cm spacing between seedlings in 2002, 2004 and 45 cm spacing in

2003.

 

'Harris Moran Seed Co., P.O.Box 4938. Modesto, CA 95352.
2Mechanical Transplanter Company, S.Central at US. 31, Holland, Michigan.

18

Application methods:

Herbicide plot size was 2.4m X 10.6m in 2002 and 2003 and 2.5m X 7.8m per
treatment in 2004. The design Of the experiment was a randomized complete block design
with four replications. The herbicides were applied as pre-transplant, post-transplant,
postemergence and post-directed applications. The list of the various herbicides applied
at different application timings are listed in Table 2. Weeded control plot was hand
weeded once in a week and continued till end of the last visual rating.

The herbicides were sprayed using a C02 backpack sprayer with a four nozzle
boom with FF8002 nozzles. Treatments were applied at the rate of 187L/ha at a pressure
Of 207 kPa and at the speed of 5.12 Kph. In case of post-directed applications, herbicides
were applied using a two nozzle shielded boom with FF11002 nozzles.

Visual rating:

Three visual ratings were recorded for all applications at regular intervals Of 7, 14,
and 21 days after the respective dates Of application. The crop and weed injury level were
scaled from 1 to 10, with one being no injury and 10 being complete death of the plant.
The visual scale was converted to percent for analysis.

Yield:

Tomato harvest began when there was visible red color on the fruit. All fruit with
red color were harvested once per week beginning in mid August and harvest continued
until frost killed the plants in late September. In 2003, both marketable and unmarketable

tomato fruit yield was recorded.

19

Statistical Analysis:

The data were subjected tO analysis Of variance by using the SAS program (SAS, 1990).
The means were separated by using Fisher’s Protected LSD at (1 =0.05 signiﬁcance level.
Because of interactions between years the results of eastern black nightshade control,

crop injury and yield are presented by year wise.

GREENHOUSE STUDY:

Greenhouse experiments were conducted twice in 2003 during the early spring
season and the mid summer season to determine the dose response Of eastern black
nightshade and tomato to sulfosulfuron and sulfentrazone.

Pre treatment of nightshade seeds:

Nightshade seeds were extracted from berries collected from the Horticulture
Teaching and Research Center (HTRC) at East Lansing, Michigan during 2002. The
nightshade seeds were subjected to cold treatment (-20 C) for 2 months and then stored in
the incubator (28 C). Study conducted by Basett and Munro (1984) Observed that addition
of 500 ppm gibberelic acid (GA3) and 0.2% potassium nitrate (KNO3) enhances
germination of black nightshade. A similar preliminary study was conducted in a growth
Chamber in 2002 to determine effects of GA; and KNO3 at different concentration on
germination of eastern black nightshade seeds. Our preliminary study observed that 1000
ppm GA; and 0.1% potassium nitrate for 10-12 hours enhanced 75% germination Of
eastern black nightshade and hence was used as a pre treatment tO trigger germination

before being sown in the ﬂats.

20

Sowing and transplanting:

Nightshade seeds were sown in ﬂats containing Baccto soil mix3. Nightshade
plants emerged within a week after sowing. During the early spring season, supplemental
light of 750 to 800 uM/mZ/s photosynthetic photon ﬂux for 14 hours per day were
provided over the ﬂats. A constant temperature of 240 C was maintained in the
greenhouse. Pikrite tomato seeds were sown at the same time in a ﬂat.

Seedlings of nightshade and tomato were transplanted at the cotyledon stage in 1
liter (L) pots ﬁlled with Baccto soil mix. During the early spring season, transplanted
seedlings were provided with supplemental lights, using high pressure sodium vapor
lamps. In mid summer season, transplanted seedlings were not provided with
supplemental light.

Treatment application:

Herbicide treatments were applied with a moving track sprayer4 with a single
80015E ﬂat-fan nozzle5 calibrated to deliver l87L/ha at a pressure Of 207 kPa. Herbicides
were applied to eastern black nightshade at the 5-6 leaf stage (5 cm in height) and to
tomato plants at 25 to 30 cm in height. Sulfosulfuron was applied at 0.0086 kg /ha,
0.0173 kg /ha, 0.034 kg /ha, 0.069 kg /ha and 0.13 kg /ha concentration. Sulfentrazone
was applied at 0.028 kg /ha, 0.056 kg /ha, 0.11 kg ai/ha, 0.22 kg /ha, and 0.44 kg /ha. The
treated nightshade and tomato seedlings were then returned to the greenhouse and

provided the respective light and temperature conditions.

 

3Baccto Soil Mix. Michigan Peat Co., P.O.Box 980129, Houston, Tx 0129.
4Allen Machine works, 607 E Miller, Midland, MI 48640.
5Teejet ﬂat fan tips. Spraying Systems Co., North Ave., Schmale Rd., Wheaton, IL 60188.

21

The treated plants were arranged in randomized complete block design with four
replications per treatment. The four replications are placed in different benches, which
were facing north-south direction.

Visual rating and harvesting of biomass:

Visual ratings were taken at 7, 14 and 21 days after treatment (DAT). Visual
ratings were scaled from 1 (no injury) to 10 (complete death) and were then converted to
percent. All aboveground plant tissue was harvested 22 DAT and fresh weights were
recorded. The plant tissues were then dried at 60 C for 6 days and their dry weights were
recorded. Herbicide effects were calculated in terms of GR50, which describes the
herbicide required for 50% dry weight reduction from untreated control.

Statistical analysis:

All the dry biomass data were subjected to ANOVA and interaction effects were
determined. The data collected from both seasons were pooled together if there were no
interactions. If there was an interaction, the data from the summer season and spring
season are presented separately.

Nonlinear regression parameters were predicted from four replications using the
polynomial quadratic model by using Sigma plot version 8 software6. This model uses the
following equation to relate dry biomass as a percentage of reduction from the control y
to the herbicide rate x.

— a+ bx+ cxz. In this 6 nation a, b, C is constant
I" Cl

 

6Sigma plot version 8.02, SPSS Inc., 233 South Wacker Drive. Chicago, IL 60606.

22

RESULTS AND DISCUSSION
FIELD STUDY:
Pre-transplant herbicide:

Pre-transplant (PRT) application Of protoporphyrinogen oxidase (PROTOX)
inhibitors like ﬂumioxazin, oxyﬂuorfen, and sulfentrazone consistently resulted in at least
95% control of eastern black nightshade in all three years (Table 2). Flumioxazin and
oxyﬂuorfen did not cause any crop injury and the effect on yield varied across the years
(Table 5). In 2002, neither ﬂumioxazin nor oxyﬂuorfen reduced yield. In 2003, lack of
rainfall during the growing season (July — August) resulted in blossom end rot, causing a
signiﬁcant reduction in yield of marketable tomatoes in all the plots. Hence there was no
difference in treatment effect on yield among the pre-transplant herbicide plots in 2003.
In 2004, ﬂumioxazin yielded 127% of the weeded control and oxyﬂuorfen yielded 75%
of the weeded control (Table 5).

PRT application of sulfentrazone controlled eastern black nightshade but caused
signiﬁcant crop injury that ranged from 18-25% in 2002 and 2004. In general, the crop
injury was relatively high during the ﬁrst week after treatment (WAT) causing necrotic
spots and curling of leaves. Sulfentrazone caused a signiﬁcant yield reduction in 2004
and yielded 53% Of the weeded control (Table 5).

Metribuzin did not control eastern black nightshade (Table 2), and did not cause
any visual injury symptoms or yield reduction in tomato in all three years when applied
PRT (Table 5). However, metribuzin controlled other broadleaf weeds in tomatoes such

as common lambsquarters and redroot pi gweed.

23

PRT application Of sulfosulfuron in 2004 resulted in 75% control of eastern black
nightshade (Table 2) and yield was 126% of the weeded control (Table 5). However in
2003, sulfosulfuron PRT did not control eastern black nightshade. We have no
explanation for the difference in level Of control between years.

Post-transplant herbicide:

There was no signiﬁcant interaction between treatments and years in 2002 and
2004 with respect to post-transplant (POT) herbicide effects on eastern black nightshade.
Sulfosulfuron gave at least 90% control Of eastern black nightshade in 2002 and 2004 but
failed to control nightshade in 2003 (Table 3). Sulfosulfuron did not cause any crop
injury and resulted in a signiﬁcant yield increase in all three years (Table 6).

Rimsulfuron, applied POT failed to control eastern black nightshade (Table 3) in
all three years and did not cause any visual injury to tomato.

Seedling shoot inhibitors such as S-metolachlor and dimethenamid-P gave 100%
control Of eastern black nightshade in all three years when applied POT (Table 3).
Activity of S-metolachlor and dimethenamid-P against eastern black nightshade was
effective from ﬁrst WAT and had a consistent prolong effect even after 6 WAT. Neither
S-metolachlor nor dimethenamid-P caused signiﬁcant crop injury in tomato in all three
years.

Napropramide was applied POT in 2002 and 2003 only. Though napropramide
belongs to the amide group and has the same mode of action as of S-metolachor and
dimethenamid-P, it did not control eastern black nightshade in 2002 or 2003 (Table 3)

and therefore napropramide was dropped from the 2004 experiment.

24

Postemergence herbicide:

Rimsulfuron and halosulfuron postemergence (POST) did not control eastern
black nightshade, nor did they injure tomato in any year (Table 4).

POST application of sulfosulfuron did not control eastern black nightshade in
2003 or 2004 (Table 4). There was no crop injury in all three years with a slight yield
reduction in 2004, resulting in yield of 87% of the weeded control (Table 7).

Sulfentrazone POST at 0.11 kg/ha and 0.22 kg/ha gave 56% and 92% control Of
eastern black nightshade respectively in 2002 (Table 4). However in 2003, sulfentrazone
0.11 kg/ha and 0.22 kg/ha gave 75% and 56% control Of eastern black nightshade
respectively, with no signiﬁcant crop injury and similar yield reduction for both rates
(Table 4,7). In 2004, there was no dose response effect from sulfentrazone and both rates
gave at least 90% control of eastern black nightshade (Table 4). Both rates Of
sulfentrazone caused signiﬁcant crop injury and yield reduction in 2004. Although it
gives fair to gOOd eastern black nightshade control, sulfentrazone probably is tOO
injurious to tomato for POST application. Typical injury symptoms were necrosis and
curling and cupping of leaves.

Metribuzin POST had no effect on eastern black nightshade and did not cause any
yield reduction or crop injury in all three years. Pyridate gave 61% and 67% control of
eastern black nightshade in 2002 and 2003, respectively (Table 4). Pyridate did not cause
any crop injury (Table 4) and yield was 83% and 85% Of the weeded control in 2002 and

2003 (Table 7).

25

Post-directed herbicides:

Carfentrazone controlled eastern black nightshade but caused unacceptable crop
injury when applied post-directed (PODIR) (Table 4). In 2002, carfentrazone water
dispersable granule (WDG) application gave 80% control of eastern black nightshade but
caused 36% crop injury and yielded 57% Of weeded control (Table 7). Carfentrazone
emulsiﬁable concentrate (BC) was used in 2003 and 2004 because the WDG was
discontinued by the manufacturer. The carfentrazone EC formulation caused 75% and
67% injury to tomato in 2003 and 2004 and resulted in yield of 67% and 22% of weeded
control (Table 7). Flumioxazin PODIR gave fair to excellent control of eastern black
nightshade in all three years. It caused no signiﬁcant yield reduction in 2002 and 2003,
but caused 39% crop injury in 2004 (Table 4). This resulted in a yield of only 13% of the
weeded control in 2004 (Table 7). Flumioxazin appears to be too toxic to tomato for

PODIR application, unless a safer method of application can be devised.

GREENHOUSE STUDY:
Dose response of sulfosulfuron:

There was a signiﬁcant difference in dose response of eastern black nightshade to
sulfosulfuron, between spring and summer season experiment (Figure 1). In the early
spring experiment, there was no dose response to sulfosulfuron. All the treatments caused
a dry weight reduction Of at least 70% from the untreated control. In the summer season
experiment there was a strong dose response (R2: 0.79) of eastern black nightshade to
sulfosulfuron, with a GRso Of 0.079 kg/ha (Table 8). In the early spring experiment, all

eastern black nightshade plants ﬂowered during the second week after treatment. This

26

may have made the plants more susceptible to the herbicide. Early ﬂowering of eastern
black nightshade might have been a response to the short days and reduced temperature
in the greenhouse in early spring. In the summer season experiment, the plants did not
ﬂower until four weeks after treatment. Even though sulfosulfuron caused reduction in
dry weight at all rates in the spring experiment, it did not give sufﬁcient control of
eastern black nightshade at 0.034 kg/ha or 0.069 kg/ha in the summer experiment.

Dose response of sulfentrazone:

The GRSO values for sulfentrazone were similar for the two seasons. GR50 values
were 0.18 kg/ha in spring and 0.14 kg/ha in summer (Table 8). There was a dose response
of eastern black nightshade to sulfentrazone and at 0.44 kg/ha dry weight reduction was
80% in spring and 95% in summer.

The results from the ﬁeld and greenhouse studies have shown that POT
application Of s-metalochlor and dimethanamid-P and PRT application of ﬂumioxazin
and oxyﬂuorfen controlled eastern black nightshade with negligible or no crop injury.
Eastern black nightshade was not controlled sufﬁciently by sulfosulfuron. Sulfentrazone

gave good eastern black nightshade control but caused unacceptable crop injury.

27

Literature cited

. Ackley, J.A., H.P. Wilson and TE. Hines. 1997. Rimsulfuron and metribuzin
efﬁcacy in transplanted tomato (Lycopersicon esculentum). Weed Technology
11:324-328.

. Bassett, I.J., and Munro DB, 1984. The biology of Canadian weeds. 67. Solanum
ptycanthum Dun., S. nigrum L. and S.sarrach0ides Sendt. Canadian Journal Of
Plant Science 65:401-414.

. Bewick, T.A., W.M. Stall, S.R. Kostewicz, and K. Smith. 1991. Alternatives for
control Of paraquat tolerant American black nightshade (Solanum americanum).
Weed Technology 5:61-65.

. Brown, J.E., W.D. Goff, W. Hogue, M.S. West, C.Stevens, V.A. Khan, B.C.
Early and LS. Brasher. 1991. Effects of plastic mulch on yield and earliness Of
tomato. Proceeding of Natural Agriculture Plastics Conference 23:21-28.

. EPA (US. Environmental Protection Agency). 1998. Methyl bromide information
In: Ozone depletion. (1998, Nov. 10), [Online] Available:
http://www.epa.gov/ozone/mbr/mbrqa.html., Dec.1 1998.

. Friesen, G.H. 1979. Weed interference in transplanted tomatoes (Lycopersicon
esculentum). Weed Science 27:11-13.

. McGiffen, Jr., ME. and J.B. Masiunas. 1991. Post-emergence control Of
broadleaf weeds in tomato. Weed Technology 5:739-745.

. McGiffen, Jr., M.E., J.B. Masiunas and J. Hesketh. 1992. Competition for light
between tomatoes and nightshades (Solanum nigrum and S. ptycanthum). Weed
Science 40:220-226.

. Milliman, L.D., D.E. Riechers, L.M. Wax and F.W. Simmons. 2003.

Characterization Of two biotypes of imidazolinone — resistant eastern black
nightshade (Solanum ptycanthum). Weed Science 51:139-144.

28

10.

11.

12.

13.

14.

15.

16.

Perez, F.G.M. and J.B. Masiunas. 1990. Eastern black nightshade (Solanum
ptycanthum) interference in processing tomato (Lycopersicon esculentum). Weed
Science 38:385-388

Reinke, H., Rosenzweig A. K.M. Clausm, C. Chisholm, and P. Jensen. 1991.
DPX-E9636, experimental sulfonylurea herbicide for potatoes. Proceeding of
Brighton Crop Protection Conference-Weeds 445-451.

[SAS] Statistical Analysis System. 1990. SAS Procedures Guides. Version 6, 3rd
edition, Cary, NC: Statistical Analysis Systems Institute 567 p.

Volenberg S., Stoltenberg E., Boerboom M. 2000. Solanum ptycanthum resistance
to acetolactate synthase inhibitors. Weed Science 481399-401

Weaver, SE. and CS. Tan. 1983. Critical period of weed interference in

transplanted tomatoes (Lycopersicon esculentum): grth analysis. Weed Science
31:467-481.

Weaver, S.E., N. Smits, and CS. Tan. 1987. Estimating yield losses of tomatoes
(Lycopersicon esculentum) caused by nightshade (Solanum spp.) interference.
Weed Science 35:163-168.

Wein, H.C. and PL. Minotti. 1987. Growth, yield, and uptake Of transplanted
fresh-market tomato as affected by plastic mulch and initial nitrogen rate. Journal
Of American Society of Horticulture Science 112: 759-763.

29

Table 1. List of treatments, application methods and the rates applied in ﬁeld experiments during 2002,
2003, and 2004 to evaluate control of eastern black nightshade in transplanted tomato.

 

 

Application day Application Application Herbicide Form Rate
method stage conc. kg ai/ha
& type
June 7, 2002 Pre-transplant Before Flumioxazin 5] WG 0.052
June 1, 2003 transplanting Metribuzin 75 DF 0.56
June 7, 2004 tomato seedlings Oxyﬂuorfen 2 L 0.28
Sulfentrazone 75 DF 0.33
Sulfosulfuron“** 75 WG 0.034
June 8, 2002 Post-transplant After transplanting Dimethenamid-P 6 EC 1.]
June 2, 2003 of tomato seedlings Napropramide“ 50 DF 2.24
June 7, 2004 Rimsulfuron 25 DF 0.034
S-metolachlor 7.62 EC 1.8
Sulfosulfuron 75 WG 0.034
July 1 l, 2002 Post -emergence Nightshades are 3- Halosulfuron“ 75 WG 0.034
July], 2003 4 leaf stage at a Metribuzin 75 DF 0.28
July], 2004 height Of 2.5-5 cm Pyridate" 3.75 EC 1.01
Rimsulfuron 25 DF 0.034
Sulfentrazone 75 DF 0.] 1
Sulfentrazone 75 DF 0.22
Sulfosulfuron“ 75 WG 0.034
July 19, 2002 Post-directed Nightshades are 3- Carfentrazone“ 40 WG 0.18
July 1, 2003 4 leaf stage at a Carfentrazone“ 2 EC 0.18
July], 2004 height Of 2.5-5 cm Flumioxazin* 5] WG 0.052

 

*Nonionic Surfactant (NIS) of 0.5% V/V was included with the respective treatment.

"”" The given treatment was applied 2002. 2003 only

*** The given treatment was applied 2003. 2004 only

30

Table 2. Pre transplant (PRT) treatment effects on eastern black nightshade (SOLPT) and crop injury rated
at six weeks aﬁer treatment in 2002, 2003 and 2004.

 

 

 

 

 

Treatments SOLPT control (%) Crop injury (%)

Rate

kg ai/ha 2002 2003 2004 2002 2003 2004
Weeded control 1003 100a 1003 0a 0 0b
Oxyﬂuorfen 0.28 100 a 100 a 100 a 6 a 0 3 b
Metribuzin 0.56 22.0 b 0 c 0 c 3 a 0 0 b
Sulfentrazone 0.33 100 a 100 a 100 a 18 b 0 25 a
Flumioxazin 0.052 94 a 100 a 94 a 1 1 a, b 0 3 b
Sulfosulfuron 0.034 NA 33 b 75 b NA 0 0 b
Lsd 17 23 8 9 NS 15

 

Values followed by the same letter in the same column are not statistically signiﬁcant at (1 =0.05.

NA - non availability of data
NS- non signiﬁcance of mean

31

Table 3. Post transplant (POT) treatment effects on eastern black nightshade (SOLPT) and crop injury rated
at six weeks after treatment in 2002, 2003 and 2004.

 

 

 

 

Treatments SOLPT control (%) Crop injury (%)

Rate

Kg ai/ha 2002 2003 2004 2002 2003 2004
Weeded control 100a 100a 100a 0 0 0
Napropramide 2.24 6 b 0 b NA 3 0 NA
S-metolachlor 1.8 100 a 100 a 100 a 11 0 0
Rimsulfuron 0.034 1 l b 0 b 0 c 0 0
Sulfosulfuron 0.034 97 a 25 c 89 b 6 0 0
Dimethenamid- P 1.1 100 a 100 a 100 a 6 0 0
Lsd 14 30 9 NS NS NS

 

 

Values followed by the same letter in the same column are not statistically signiﬁcant at (1 =0.05
NA — Treatment not included in that respective year
NS- non signiﬁcance of mean

32

Table 4. post emergence (POST) and post directed (PODIR) treatment effects on eastern black nightshade
(SOLPT) and crop injury rated at three weeks after treatment in 2002, 2003 and 2004.

 

 

 

 

Treatments SOLPT control (%) Crop injury (%)

Rate

Kg ai/ha 2002 2003 2004 2002 2003 2004
POST
Weeded control 100a 100a 100a 0 0a
Rimsulfuron 0.034 6 0 C 25 b 3 0 0 a
Sulfosulfuron" 0.034 67 a, b 6 c 1 1 b 7 0 0 a
Halosulfuron" 0.034 11 c 0 c 0 b 3 11 0 a
Metribuzin 0.28 1 1 c 0 c 0 b 3 0 0 a
Sulfentrazone 0.11 56 b 75 b 94 a 1 1 0 11 b
Sulfentrazone 0.22 92 a 56 b 100 a 8 7 22 c
Pyridate 1.01 61 b 67 b NA 0 0 NA
Lsd 31 41 29 NS NS 10
PODIR
Weeded control 100 100 100 0b 0b 0c
Carfentrazone“ 0.18 83 100 100 36 a 75 a 67 a
Flumioxazin* 0.052 64 100 100 6 b 14 b 39 b
Lsd NS NS NS 1 1 16 6

 

 

Values followed by the same letter in the same column in each application method are not statistically

signiﬁcant at (1 =0.05.

*Nonionic Surfactant of 0.5% V/V was included with herbicide
NA - Treatment not included in that respective year

NS- non signiﬁcance of mean

33

Table 5. Effect of pre-transplant (PRT) treatments on tomato yield from 2002-2004.

 

 

 

Yield ( kg/plot) Yield as % of weeded control

Treatments Rate

kg ai/ha 2002 2003 2004 2002 2003 2004
Weeded control 146 39 c 85 a, b 100 100 100
Oxyﬂuorfen 0.28 125 61 a, b 63 b, c 86 156 75
Metribuzin 0.56 148 55 a. b 86 b, a 101 141 101
Sulfentrazone 0.33 124 70 a 45 c 85 179 53
Flumioxazin 0.052 122 67 a 108 a 84 172 127
Sulfosulfuron 0.034 NA 52 a. b 107 a NA 133 126
Lsd 37 23 30

 

 

Values followed by the same letter in the same column are not statistically signiﬁcant at 0. =0.05

NA — Treatment not included in that respective year

NS- non signiﬁcance of mean

34

Table 6. Effect of post-transplant (POT) treatments on tomato yield from 2002-2004.

 

 

 

 

Yield ( kg/plot) Yield as % of weeded control

Treatments

Rate

2002 2003 2004 2002 2003 2004

Kg ai/ha '
Weeded control 146 a,b 39 85 100 100 100
Napropramide 2.24 131 a, b 58 NA 90 149 NA
S-metolachlor 1.8 123 b 39 98 84 100 1 15
Rimsulfuron 0.034 132 a, b 40 114 90 103 134
Sulfosulfuron 0.034 169 a 54 120 1 16 138 141
Dimethenamid-P 1.1 152 a, b 60 86 104 154 101
Lsd 45 NS NS

 

 

Values followed by the same letter in the same column are not statistically signiﬁcant at 0. =0.05
NA- Treatment not included in that respective year
NS- non signiﬁcance of mean

35

Table 7. Effect of post emergence (POST) and post directed (PODIR) treatments on tomato yield from
2002-2004.

 

 

 

Treatments Yield ( kg/plot) Yield as % of weeded control
Rate
Kg ai/ha 2002 2003 2004 2002 2003 2004
POST
Weeded control 146 a 39 a. b 85 a 100 100 100
Rimsulfuron 0.034 140 a,b 30 b 72 ab 96 77 85
Sulfosulfuron" 0.034 138 a,b 51 a,b 74 a,b 95 131 87
Halosulfuron* 0.034 123a,b, c 47 a, b 68 be 84 121 80
Metribuzin 0.28 138 a, b 52 a. b 90 a 95 133 106
Sulfentrazone 0.1 1 88 c 37 a. b 39 c 60 95 46
Sulfentrazone 0.22 102 b. c 62 a 40 c 70 159 47
Pyridate 1.01 121a,b,c 33 b NA 83 85 NA
Lsd 40 27 35
PODIR
Weeded control 146 a 39 85 a 100 100 100
Carfentrazone* 0.18 83 b 26 19 b 57 67 22
Flumioxazin“ 0.052 120 a. b 54 1 l b 82 138 13
Lsd 43 NS 22

 

 

Values followed by the same letter in the same column in each application method are not statistically

signiﬁcant at (1 =0.05.

*Nonionic Surfactant of 0.5% V/V was included with herbicide
NA - Treatment not included in that respective year

NS- non signiﬁcance of mean

36

Table 8. Polynomial quadratic equations, R2 values*, GR50 values to determine the response of eastern
black nightshade (SOLPT) and tomato to sulfosulfuron", sulfentrazone treatments applied during two

different season.

 

 

Treatment Time of Leaf r2 y= a+bx+c x2 GRSO
application stage/ (kg/ha)
height
SulfosulfuronM SOLPT-Early 6 Leaf/ 0.038 y= 71.08-79.29x-677.20 x2 <0.0086
Spring 5 cm
SOLPT-Mid 6 Leaf/ 0.79 y= -5.37+758.89x-805.84 x2 0.079
summer 5 cm
Tomato*** 25-30 0.21 y= -4.44+442.56x-1969.71 x2 >0.13
cm
Sulfentrazone SOLPT-Early 6 Leaf/ 0.85 y= -17.50+472.51x-583.78 x2 0.18
Spring 5 cm
SOLPT-Mid 6 Leaf/ 0.82 y= -17.36+567.59x-712.38x2 0.14
summer 5 cm
Tomato*** 25-30 0.32 y=-11.11+418.01x-658.58 x2 0.22
cm

 

 

 

 

* All R2 values are signiﬁcant at a < 0.01
“Nonionic Surfactant of 0.5% V/V was included with herbicide
*** Interactions between two seasons are not signiﬁcant and the data are pooled together

37

If

100

 

 

 

 

80-1
S3
c 601
.9
‘6
3
"C
93 40‘
E
.9
G)
3 20-
2‘
D
0-
'20 I I I I I I r

 

0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 0.16

Sulfosulfuron ( kg ai/ha)

 

Mid summer SOLPT response (R2 = 0.79)
_ .— —- Early spring SOLPT response ( R2 = 0.19)
------------- Tomato response (R2 =O.42)

 

 

 

Figure 1. Polynomial quadratic comparison of eastern black nightshade (SOLPT)
response to sulfosulfuron at 0.0086 kg/ha, 0.0173 kg/ha, 0.034 kg/ha, 0.069 kg/ha and
0.13 kg/ha concentrations applied during early 2003 spring and mid summer season in

greenhouse.

38

 

 

 

 

 

100
80 -
Zs‘
: 60 -
.9
*6
3
‘O
93 4o -
.‘c"
.9
d)
3 20 J
2‘
o
0 a
'20 I I I I
0.0 0.1 0.2 0.3 0.4 0.5

Sulfentrazone ( kg ai/ha)

 

— — — Early spring SOLPT response (R2 =O.85 )
Mid summer SOLPT response (R2 = 0.82 )
------------- Tomato response (R2 = 0.32 )

 

 

 

Figure 2. Polynomial quadratic comparison Of eastern black nightshade (SOLPT)
response to sulfentrazone at 0.028 kg /ha, 0.056 kg /ha, 0.11 kg ai/ha, 0.22 kg /ha, and
0.44 kg fha concentrations applied during early 2003 spring and mid summer season in

greenhouse.

39

CHAPTER 3
VARIATION IN HERBICIDE RESPONSE AMONG NIGHTSHADE (Solanum
spp) AND GROUNDCHERRY (Physalis spp) SPECIES AND WITHIN
NIGHTSHADE POPULATIONS

Abstract. Greenhouse experiments were conducted in 2003 and 2004 to determine the
variation in herbicide response among nightshade and groundcherry species and within
nightshade populations in Michigan. Eastern black nightshade populations had a wider
range of variation than hairy nightshade and horsenettle population to metribuzin,
sulfentrazone and halosulfuron. Among the 12 eastern black nightshade populations
screened for herbicide response, the Ingham2 population had greater tolerance to
herbicides than the other populations. A wide range of variation was noted among hairy
nightshade populations in response to halosulfuron. Among the eight populations of hairy
nightshade, the Macombl and Bay2 population was more tolerant than the other
populations to halosulfuron. There was no signiﬁcant difference in herbicide tolerance
among horsenettle populations. Among nightshade and groundcherry species, eastern
black nightshade was the only species that had a high range of tolerance to metribuzin.
Clammy groundcherry was more tolerant to sulfentrazone than eastern black nightshade,
hairy nightshade, horsenettle and smooth groundcherry. Smooth groundcherry was more
tolerant to pyridate than eastern black nightshade, hairy nightshade, horsenettle and
clammy groundcherry.

Key words: Eastern black nightshade (Solanum ptycanthum), hairy nightshade (Solanum
sarrachoides), horsenettle (Solanum carolinense), smooth groundcherry (Physalis

subglabrata), clammy groundcherry (Physalis heterophylla)

40

 

 

INTRODUCTION

Weeds in the nightshade family (Solanaceae) are difﬁcult to control in vegetable
cropping systems throughout the northeastern United States. Solanaceous weeds such as
eastern black nightshade (Solanum ptycanthum), hairy nightshade (Solanum
sarrachoides) and horsenettle (Solanum carolinense) are more troublesome in tomato,
potato and pepper than clammy groundcherry (Physalis heterophylla) and smooth
groundcherry (Physalis subglabrata). Nightshade causes direct yield losses due to plant
competition with the crop and indirect yield losses by hosting a number Of pests of
Solanaceous crops. Perez and Masiunas (1990) estimated that four eastern black
nightshade plants per square meter reduced yield by 60% in transplanted tomato by
interfering with tomato growth within 4 to 8 weeks after crop establishment. In potato,
hairy nightshade acts as an alternate host to potato leaf roll virus (Thomas, 2002) and
Rhizoctonia solani (Ogg, 1989) and contributes to the transmission Of diseases from one
season to the next, leading to indirect yield reduction of potato. Similarly, horsenettle acts
as an important reservoir Of Colorado potato beetle (Covarrubias et al., 1996) causing
indirect yield reduction in potato.

Since nightshade and tomato belong to the same botanical family, nightshades and
tomato have a close genetic relationship and share similar growth habitats, physiology
and herbicide susceptibilities (McGiffen & Masiunas, 1991; Perez & Masiunas, 1990;
Weaver et al., 1987). Hence nightshades are difﬁcult to control in tomato. In addition,
there are few registered postemergence herbicides for nightshade control in tomatoes.

The similarities Of closely related nightshade species has led to improper

identiﬁcation Of nightshades in the past and has caused conﬁision with regard to the

41

response of nightshades to herbicides. The revision of taxonomic descriptions by
Schilling (1981), reports on nightshade species variation in response to herbicides (Ogg,
1986) and the difference in germination response to differing cultural practices (Ogg et
al., 1981; Holm et al., 1977) has lead to accurate identiﬁcation of nightshade species.

Differences in eastern black nightshade and hairy nightshade response to
aciﬂuorfen (Majek, 1981), chlorsulfuron (Ogg et al., 1981) and rimsulfuron (Ackley,
1997) have been reported. Recently eastern black nightshade biotypes resistant to ALS
inhibitors have been reported in Wisconsin (Volenberg et al., 2000), Indiana and Illinois
(Milliman et al., 2003). Furthermore, there are reports substantiating herbicide variation
among accessions Of eastern black nightshade (Ogg, 1986). There are no reports in
variation in herbicide response among hairy nightshade and horsenettle populations.
Richman et a1. (1995) reported phylogenetic differences in two populations of horsenettle
but there was no report on herbicide sensitivity in horsenettle populations.

The Objective of this study was to determine any variation in sensitivity among
nightshade and groundcherry species to herbicides and also to determine variation in

herbicide sensitivity within the populations Of nightshade species.

42

 

MATERIALS AND METHODS
Greenhouse experiments were conducted in 2003 and 2004 to evaluate the herbicide dose
response among eastern black nightshade, hairy nightshade, horsenettle, smooth
groundcherry and clammy groundcherry populations collected in Michigan.
Collection of berries:

In 2002, 1] populations of eastern black nightshade, seven populations Of hairy
nightshade, four populations of horsenettle and one population each of clammy
groundcherry and smooth groundcherry were collected from locations in Michigan.
Location, soil type, associated crops, and dates of collection of the weed populations are
listed in Tables 1-4. The berries Of ten nightshade plants in each ﬁeld were randomly
collected and bagged together.

Processing of berries:

The berries of these weeds were crushed and soaked in water for one night. Fruit
juices were removed by wrapping in cheese cloth and squeezing by hand. Then the seeds
in the Cheese cloth were washed repeatedly and air dried for a week. The seeds were
stored for two months at -20°C and then stored in incubator at 28° C to overcome
dormancy.

Pre treatment of seeds:

A preliminary study was conducted in a growth chamber in 2002 to determine
effects of germination enhancing chemicals on these seeds. Soaking seed in Gibberrellic
acid (GA3) at 1000 ppm and 0.1% potassium nitrate (KnO3) for 10-12 hours resulted in at

least 75% germination. Therefore seeds were pretreated overnight with 1000 ppm GA3

43

 

 

and 0.1% KN03 to enhance germination. The pretreated seeds were allowed to air dry
and then were sown in the ﬂats containing Bacctol soil mix.
Sowing and transplanting:

Seeds were sown during the ﬁrst week of June in the greenhouse and transferred
to 1L pots, when the seedlings had 2 tO 3 true leaves. The seedlings were maintained in
the pots until the four to six leaf stages, and then were subjected to postemergence
herbicide treatments.

Treatments and application:

The herbicide treatments and concentrations are listed in Table 5. The 1 X rates
for the herbicide treatments in Table 5 were used in the ﬁeld experiment in 2002, 2003
and 2004. All the weed populations were Sprayed in three consecutive days, by using a
single tip track sprayer3 with an 80015E ﬂat-fan nozzle2 calibrated to deliver 187 L/ha at
a pressure of 207 kPa. The treated nightshade populations were placed in the greenhouse
and were arranged in randomized complete block design with four replications per
treatment. The four replications are placed in different benches, which were facing north-
south direction.

Visual rating and biomass measurements:

Visual ratings were taken at 1, 2 and 3 weeks after treatment (WAT). Visual ratings were
scaled from 1 (no injury) to 10 (complete death) and then converted to a percentage for
statistical analysis. Plants were harvested by clipping at the stem base region at 3 WAT

and the fresh weights were recorded. The plants were air dried at 60°C for a week and

 

IBaccto Soil Mix. Michigan Peat Co., P.O.Box 980129, Houston, TX 0129.
2Teejet ﬂat fan tips. Spraying Systems Co., North Ave., Schmale Rd., Wheaton, IL 60188.
3Allen Machine works, 607 E Miller, Midland, MI 48640.

44

 

then weighed again. The plant weights were then converted to percent of reduction from
the untreated control.
Statistical analysis:

The two years data were subjected to ANOVA to determine the signiﬁcance Of
year x year interactions and data were combined together over years if there was no year
effect. Using a log-logistic model (Seefeldt et al., 1995), a non linear regression curve
was developed by using Table curve4 and Sigma plot version 8 softwares. The following
log-logistic equation, relates the percent Of dry biomass reduction from the control y to
the herbicide rate x (Seefeldt et al., 1995).

y=C+ D—C

 

1+ (x/GR 50) b
In this equation, C is the lower response limit, D is the upper response limit, b is the slope

and GRSO is the rate which resulted in a 50% reduction in biomass.

 

4Jandel Scientiﬁc, Table Curve 2D v. 3.1, 2591 Kemer Boulevard, San Rafael, CA 9490].
5Version 8.02, SPSS Inc., 233 South Wacker Drive, Chicago, IL 60606.

45

 

 

RESULTS AND DISCUSSION
Response of eastern black nightshade populations to herbicides:

Eastern black nightshade populations had a wide range Of variation in response to
metribuzin at rates higher than 0.28 kg/ha. Dose response curves for the 12 populations in
response to metribuzin are presented in Figures 1a, 1b, and 1c. Among the 12
populations, the Newaygo2 and Monroe4 populations were the most susceptible to
metribuzin with a 50% growth reduction (GR50) at 0.31 kg/ha (Figurela). Oceanal,
Inghaml, Ingham2, Bay] and Monroe2 populations had high tolerance to metribuzin at
1.12 kg/ha and could not ﬁt the log logistic equation (Table 6). In general, all eastern
black nightshade populations were tolerant tO metribuzin at the normal dose rate Of 0.28
kg/ha.

Sulfentrazone, a protoporphyrinogen oxidase inhibitor, reduced the dry weight of
all eastern black nightshade populations. However, populations varied in their
susceptibility to sulfentrazone (Figures 2a, 2b, 2c, 2d). Newag02 was the most
susceptible among the eastern black nightshade populations with a GRSO of 0.036 kg/ha
(Table 7). Inghaml & Ingham2 populations were the most tolerant Of sulfentrazone
(Figure 2c) with a GRSO =0.22 kg/ha which is twice the standard rate. The Ingham2 R/S
(Resistance/ Susceptible) ratio was calculated at about 6 times more tolerant than the
Newag02 (Figure2d). Eastern black nightshade populations within counties other than
Monroe had similar responses. Among Monroe populations, Monroe] was 60% more
tolerant than Monroe2 (Table 7). Log-logistic equations could not be Obtained for the
monroe4 population response to sulfentrazone due to high degree Of non homogenous

data.

46

Eastern black nightshade populations other than Macomb3 and Macombl were
tolerant to halosulfuron. However, there was a wide range of variation among the
populations above the rate of 0.034 kg/ha except in Macomb3 and Macombl populations
(Figures 3a, 3b, 3c). Macomb3 and Macombl populations were the most susceptible to
halosulfuron with a GRso less than the normal rate of 0.034 kg/ha (Table 8). In contrast,
the Macomb2 population was 6 times more tolerant to halosulfuron than Macomb3, and 4
times more tolerant than the Macombl population (Figure 3c). The Ingham2 population
was very highly tolerant to halosulfuron and could not ﬁt the log logistic equation (Table
8). A GR50 value could not be obtained for other populations such as Bay], Inghaml,
Monroe2, Mason] and Newag02 due to high degree of tolerance to halosulfuron.

There was a wide range of variation among eastern black nightshade populations
to sulfosulfuron (Figures 4a, 4b, 4c). The Monroe2 population was highly susceptible to
sulfosulfuron, with a GRSO of 0.032 kg/ha. The Oceanal population was the most tolerant
to sulfosulfuron among the eastern black nightshade populations (Figure 4d). The GR50
value for the Oceanal population could not be calculated and also had a low R square
value due to high tolerance (Table 9). The Macomb3 population possessed GR50 Of 0.10
kg/ha which was two times more tolerant than the highly susceptible Macomb2
population.

Most Of the populations Of eastern black nightshade had a high degree Of
sensitivity to pyridate (Table 10) and there was no wide variation among eastern black
nightshade populations susceptibility to pyridate (Figure 5a). Ingham2 and Oceanal were
susceptible to pyridate at 0.44 kg/ha and 0.36 kg/ha respectively (Table 10). Mason],

Inghaml, and Bay] populations were susceptible at 0.29 kg/ha. Other eastern black

47

nightshade populations were highly sensitive to pyridate even at the very low rate Of
0.225 kg/ha and hence log-logistic equation could not be applied.

In conclusion, the Ingham2 population had greater tolerance to all herbicides than
the other eastern black nightshade populations. The Oceanal population was more
tolerant to sulfosulﬁrron and metribuzin than other eastern black nightshade populations.
Past cultural practices by using herbicides in Ingham2 and Oceana] for many years may
be the reason for greater tolerance to herbicides.

Response of hairy nightshade populations to herbicides:

In general, hairy nightshade populations were highly susceptible to metribuzin
(Figure 5b, 5C). The Montcalm] population was the most susceptible among the 7
populations with a GR50 of 0.10 kg/ha (Table 11). The Macombl population was twice as
tolerant compared to the Montcalm] population with a GRSO of 0.23 kg/ha (Figure 5d).

Most of the hairy nightshade populations were highly susceptible to sulfentrazone
(Figure 6a, 6b, Table 12). The Presque2 and Presque3 populations were the most
susceptible among the hairy nightshade populations with a GR50 less than 0.028 kg/ha.
The Macomb] population was the most tolerant to sulfentrazone compared to other hairy
nightshade populations with a GR50 Of 0.10 kg/ha, and was four times more tolerant than
the Presque2 and Presque3 populations (Figure 6c).

Hairy nightshade populations were highly susceptible sulfosulfuron (Figures 7a,
7b) and the range of susceptibility varied from <0.0086 kg/ha to 0.037 kg/ha. Log-
logistic equations could not be obtained for the Presquel and Montcalm2 populations due
to a high degree of susceptibility to sulfosulfuron. In general, the Presque2, Presque3, and

Presque4 populations were similar in dose response to sulfosulfuron (Table13) with a

48

GR50 of 0.0099 kg/ha. The Macomb] and Montcalm] populations were more tolerant to
sulfosulfuron compared to other hairy nightshade populations with a GRSO of 0.037 kg/ha.
The Macombl and Montcalm] populations were 4 times more tolerant to sulfosulfuron
than the Presque2, Presque3, Presque4 populations (Figure 7c).

Hairy nightshade populations had a wide range Of variation in response to
halosulfuron (Figure 8a). The Presque2 population was the most susceptible among the
hairy nightshade populations with GRsn Of 0.021 kg/ha. The Macombl and Bay2
populations were tolerant to halosulfuron. Due to the high degree Of tolerance, GR50
values could not be calculated (Figure 8b). The Presque4 population (GR50 = 0.077 kg/ha)
was three times more tolerant than the Presque2 population (Table14). Presquel and
Montcalm] populations could not be analyzed for response to halosulfuron, due to the
high degree of non homogenous variability in the data which could not be transformed.

All hairy nightshade populations were sensitive to pyridate at 0.225 kg/ha and
data could not be ﬁtted to a log logistic equation (Table 15).

Among all hairy nightshade populations, the Macombl population was more
tolerant than the other hairy nightshade populations in response to all of the herbicides.
Response of horsenettle populations to herbicides:

Horsenettle populations were very susceptibile to halosulfuron (Figure 9). Among
the four populations, Monroe5 was highly susceptible and a GR50 value could not be
Obtained. Besides the Monroe5 population, Berrien2 was very sensitive with GR50 of
0.0086 kg/ha (Table16).

Narrow range Of variation was Observed among horsenettle populations in

response to sulfentrazone (FigurelO). Among the four horsenettle populations,

49

Montcalm3 was the most susceptible to sulfentrazone with GR50 Of 0.012 kg/ha.
Vanburenl was the most tolerant among the four populations with a GRso Of 0.35 kg/ha
which was three times more tolerant than Berrien2 and 30 times more tolerant than
Montcalm3 populations in response to sulfentrazone (Table 17).

Horsenettle populations were highly susceptible to sulfosulfuron. A narrow range
of variation was observed among the four horsenettle populations in response to
sulfosulfuron. (Table 18). Vanburenl was the most susceptible to sulfosulfurOn and had
GR50 less that 0.0086 kg/ha. All horsenettle populations were very sensitive below the
normal dose of 0.034 kg/ha (Figure 11).

Horsenettle populations had a narrow range of variation in response to pyridate.
All Of the horsenettle populations were sensitive to pyridate at rate less than 0.5 kg/ha
(Figurel2). The Monroe5 population was the most sensitive among the horsenettle
populations, (Table 19) with a GR50 less than 0.252 kg/ha.

All horsenettle populations also had a high degree of sensitivity to metribuzin
even at the lowest rate and hence the log logistic equation could not be developed (Table
20).

In general, there was not much variation in tolerance among the four horsenettle
populations to herbicides.

Response of clammy groundcherry to herbicides:

The Newagol population had a high tolerance to sulfentrazone and a GR50 could
not be obtained at the given rates (Table 21). The low susceptibility of Newagol
population to sulfentrazone might be due to the presence Of dense pubescence on the

leaves that might have prevented the entry Of sulfentrazone into the cuticle of the leaves.

50

 

However, the Newagol population was sensitive to sulfosulfuron, metribuzin and
pyridate. Due to the high degree Of non homogenous variability in data with respect to the
halosulfuron treatment, a log logistic curve could not be applied.

Response of smooth groundcherry to herbicides:

The Ingham3 population was very sensitive to sulfentrazone treatment with a
GR50 of 0.104 kg/ha. There was a high degree of sensitivity to metribuzin in the Ingham3
population and log logistic curve could not be developed due to a high degree of
sensitivity (Table 22). The Ingham3 population was also susceptible to halosulfuron at
0.038 kg/ha and pyridate at 1.39 kg/ha (Table 22). With respect to sulfosulfuron,
Ingham3 possessed fair tolerance, with a GRsn of 0.091 kg/ha.

The results of this study indicate that nightshade and groundcherry species
respond differently to a wide variety Of herbicides (Table 23). Past cultural practices in
using herbicides for many years might be the reason for higher tolerance among eastern
black nightshade populations and some hairy nightshade populations to herbicides.
Eastern black nightshade populations in particular have a wide range of variation to
herbicides (Table 23). Eastern black nightshade populations such as Ingham2 and
Oceanal may possess multiple resistant characteristics to triazine and sulfonylurea
herbicides and might be likely to build resistance to other herbicides. Studies conducted
by Vollenberg et al., (2000) and Milliman et al., (2003) have reported ALS resistant
eastern black nightshade in agronomic crops. These results indicate that eastern black
nightshade may become an increasingly troublesome weed among Solanaceous species in

tomato and likely to build resistance to many herbicides.

51

Literature cited

. Ackley, J.A., H.P. Wilson and TE. Hines. 1997. Rimsulfuron and metribuzin
efﬁcacy in transplanted tomato (Lycopersicon esculentum). Weed Technology
112324-328.

. Covarrubias M.J., F .A. Drummond, and D.L. Haynes. 1996. Population dynamics
Of the Colorado potato beetle on horsenettle in Michigan. Entomological Society
of America 25:68-77.

. Holm, L.G., D.L. Pluckett, J.V. Pancho and JP Herberger. 1977. The Worlds

Worst Weeds. Solanum nigrum — Distribution and Biology. University Of Hawaii
Press. 403-434.

. Majek, BA. 1981. Nightshade control in ﬁeld crops. University of Minnesota
Agricultural Extension folder 60322-4.

. McGiffen, Jr., ME. and J.B. Masiunas. 1991. Post-emergence control of
broadleaf weeds in tomato. Weed Technology 5:739-745.

. Milliman, L.D., D.E. Riechers, L.M. Wax and F.W. Simmons. 2003.

Characterization of two biotypes of imidazolinone -— resistant eastern black
nightshade (Solanum ptycanthum). Weed Science 51: 139-144.

. Ogg A.G., Jr., B.S. Rogersand and BE. Schilling. 1981. Characterization of black

nightshade (Solanum nigrum) and related species in the United States. Weed
Science 29227-32.

. Ogg A.G., Jr. 1986. Variation in response Of four nightshades (Solanum spp.) to
herbicides. Weed Science 34:765-772.

. Ogg, A.G., Jr. 1989. Taxonomy, distribution, biology and control of black
nightshade (Solanum nigrum) and related species in the United States and Canada.
Weed Science 4:25-58.

52

 

10.

11.

12.

13.

14.

15.

16.

17.

Perez, F .G.M. and J.B. Masiunas. 1990. Eastern black nightshade (Solanum
ptycanthum) interference in processing tomato (Lycopersicon esculentum). Weed
Science 382385-388

Richman, A.D., T.H. Kao, S.W. Schaeffer, and MK. Uyenoyama. 1995. S-allele
sequence diversity in natural populations of Solanum carolinense (Horsenettle).
Heredity 75:405-415.

Schilling, BE. 1981. Systematics of Solanum sect. Solanum (Solanaceae) in
North America Systematic Botany 6: 1 72-185.

Seefeldt, S.S., J.E. Jensen and ER Fuerst. 1995. Log-logistic analysis of
herbicide dose- response relationships. Weed Technology 9:218-227.

Thomas, PE. 2002. First report of Solanum sarrachoides (hairy nightshade) as an
important host of potato leafroll virus. Plant Disease 86:559.

Volenberg S., E. Stoltenberg and M. Boerboom. 2000. Solanum plycanthum
resistance to acetolactate synthase inhibitors. Weed Science 48:399-40]

Weaver, SE. and CS. Tan. 1983. Critical period of weed interference in

transplanted tomatoes (Lycopersicon esculentum) growth analysis. Weed Science
31:467-481.

Weaver, S.E., N. Smits and CS. Tan. 1987. Estimating yield losses Of tomatoes
(Lycopersicon esculentum) caused by nightshade (Solanum spp.) interference.
Weed Science 35:163-168.

53

 

Table 1. Eastern black nightshade (SOLPT) populations collected from various regions in
Michigan; including location, current crop and soil properties.

 

Soil Properties

 

 

SOLPT Location Current
Population ( Michigan) Crop Soil pH Organic N(NO3)
Type Content (%) (ppm)
Newag02 T12N, R14W Tomato Loamy 5.7 2.98 18.6
Sec : 17, sand
Sheridan twp
Inghaml T3N, R1W Tomato Sandy 7.1 2.14 5.7
Sec : 12, loam
Delhi twp
Ingham2 TIN, R2E Potato organic 4.7 32.41 45.3
Sec : 35,
Stockbridge twp
Oceanal T15N, R17W Asparagus Loamy 5.9 1.91 9.]
Sec : 32, sand
Hart twp
Masonl T18N, R17W Snap bean Sandy 7 2.62 51.]
Sec : 29 clay
Riverton twp loam
Monroe2* T7S, R6E Tomato - - - -
Sec : 21,
Dundee twp
Monroe4 T7S, R6E Pepper Clay 6.8 2.98 5.8
See : 21, loam
Dundee twp
Macombl T3N, R13E Tomato Sandy 5.7 1.52 l 1.]
Sec : 17, loam
Macomb twp
Macomb2 T4N, R13E Tomato Sandy 7 3.57 17.2
Sec : 5, clay
Ray twp Loam
Macomb3 TSN, R13E Tomato Loamy 5.3 2.34 32.9
See : 32, sand
Armada twp
Bayl** T15N, R3E Potato - 7.3 2.21 14.2
Sec : 22,
Garﬁeld twp

 

 

*Soil properties of the given population is not available
** Soil type of the given population is not available

54

Table 2. Hairy nightshade (SOLSA) populations collected from various regions in

Michigan; including location, current crop and soil properties.

 

Soil Properties

 

 

SOLSA Location Current
Population ( Michigan) Crop Soil Organic N( N03)
Type pH Content (ppm)
(”/0
Montcalm] T12N, R6W Potato Sandy 6.2 3.1 14.6
Sec : 34, loam
Home twp
Montcalm2* T11N,R9W A' Sandy 6.1 1.17 -
Sec : 25. loam
Maple valley twp
Macombl T3N, R13E Tomato Sandy 5.7 1.52 11.]
Sec : 17, loam
Macomb twp
Presquel T34N, R5E Potato Sandy 7.6 1.4 15.9
Sec : 2, loam
Belknap twp
Presque2 T34N, R6E Dry bean Loam 7 3.03 8.5
Sec : 30,
S.Pulawski twp
Presque3 T34N, R5E Dry bean Sandy 7.8 3.43 6.]
Sec : 21, clay
Belknap twp loam
Presque4** T34N, R6E Dry bean - - - -
Sec : 30,
S.Pulawski twp
Bay2 T15N, R3E Potato Sandy 7.3 2.21 14.2
Sec : 22, loam
Garﬁeld twp

 

 

* Nitrogen content of the given population is not available

** Soil properties Of the given population is not available

A1 The given population is collected frOm agrestal region (uncultivated/ undisturbed

region)

55

Table 3. Horsenettle (SOLCA) populations collected from various regions in Michigan;

including location, current crop and soil properties.

 

 

 

SOLCA Location Current. Soil Properties
Population ( Michigan) Crop
Soil Organic
type pH Content N (NO3)
(%) (Ppm)
Berrien2 T5S, R18W A' Sandy 6.2 1.71 43
Sec : 15, loam
Sodus twp
Vanburenl T3S, R16W Tomato Sandy 6.5 1.71 14.3
Sec : 1, loam
Hartford twp
Montcalm3 T10N, R8W Snapbean Sandy 5.4 2.07 14.8
Sec : 5 loam
Monroe5* T7S, R6E Soybean
Sec : 30,
Dundee twp

 

 

AI The given population is collected from agrestal region (uncultivated/ undisturbed
region

*SOil properties Of the given population is not available

56

Table 4. Clammy groundcherry (PHYHE) and smooth groundcherry (PHYSU)

populations collected from various regions in Michigan; including location, current crop

and soil properties.

 

 

 

Species Location Assoc. Soil Properties
( Michigan) Crop
Soil pH Organic N(NO3)
ije Content (%) (ppm)
PHYHE T12N, R14W Tomato Loamy 6.2 2.05 8.6
(Newagol) Sec : 36, sand
Sheridan twp
PHYSU T2N, R2W Onion organic 6.6 71.77 102.1
(Ingham3) Sec : l4,
Aurelius twp

 

 

57

 

Table 5. Treatments applied at various concentrations to eastern black nightshade, hairy

nightshade, horsenettle, smooth groundcherry and clammy groundcherry populations.

 

 

Treatments 0.25X 0.5X 1X 2X ' 4X
(kg ai/ha) (kg ai/ha) (kg ai/ha) (kg ai/ha) (kg ai/ha)
Metribuzin 0.070 O. 14 0.28 0.56 1.12
Sulfentrazone 0.028 0.056 0.1 12 0.224 0.448
Sulfosulfuron 0.0086 0.0173 0.0347 0.0694 0.1388
Halosulfuron 0.0086 0.0173 0.0347 0.0694 0.1388
Pyridate 0.252 0.50 1.01 2.02 4.04

 

 

 

 

hm A: 4! us

Table 6. Log logistic equations, R2 values and GR50 values for eastern black nightshade

(SOLPT) populations in response to metribuzin treatment.

 

 

SOLPT R 2 a Log-logistic equation GR 50( kg ai/ha)
Population

Monroe4 0.78 y= 0 + 74.86/[1+(x/0.31)"3~°‘] 0.31
Newayg02 0.53 y= 0+ 76.89/[1+(x/0.31)"'76] 0.31

Masonl 0.66 y= 0+ 86.71/[1+(x/0.50)'O‘74] 0.50
Monroe] 0.82 y= O+84.85/[1+(x/0.60)"9'49] 0.60
Macomb2 0.61 y= 0+93.79/[1+(x/0.61)"951 0.61
Macomb] 0.74 y= 0+87.19/[1+(x/0.67)""0] 0.67
Macomb3 0.72 y= 0+70.80/[1+(x/0.82)"-37] 0.82
Oceanalb >1.12
Inghaml *’ >1.12
Ingham2 b >1.12

Bay] b >1.12

Monroe2 b >1 .12

 

a All R2 values are signiﬁcant at 01 < 0.01.

b Populations were highly tolerant and could not ﬁt in to the equation and have low R2.

59

 

 

Table 7. Log logistic equations, R2 values and GR50 values for eastern black nightshade

(SOLPT) populations in response to sulfentrazone treatment.

 

 

SOLPT R 23 Log-logistic equation GRsn
Population (kg ai/ha)
Newag02 0.36 y= 0+88.11/[1+(x/0.036)'°‘33] 0.036
Macombl 0.90 y= 0+ 102.84/[1+(x/0.066)"~7°] 0.066
Macomb2 0.76 y= 0+ 95.09/[1+(x/0.07)"-"4] 0.07
Macomb3 0.78 y= 0+ 86.68/[1+(x/0.07)“-6'] 0.07
Monroe2 0.45 y= 0+80.18/[1+(x/0.091)'°°35] 0.091
Oceanal 0.75 y= O+79.60/[1+(x/0.091)'5‘5'] 0.091
Masonl 0.91 y= 0+ 94.01/[1+(x/0.12)'3'36] 0.12
Bay] 0.82 y= 0+101.69/[1+(x/0.14)’3'09] 0.14
Monroe] 0.79 y= 0+ 98.46/[1+(x/0.15)'°-74] 0.15
Ingham2 0.67 y== 0+95.76/[1+(x/0.22)‘19‘89] 0.22
Inghaml 0.65 y= 0+ 81.17/[1+(x/0.22)'0‘74] 0.22
Monroe4 b

 

 

a All R2 values are signiﬁcant at 01< 0.01

b Population has huge variability in data which could not be transformed for analysis

60

Table 8. Log logistic equations, R2 values and GR50 values for eastern black nightshade

(SOLPT) populations in response to halosulfuron treatment.

 

 

SOLPT R 2" Log-logistic equation OR50
Population ( kg ai/ha)
Macomb3 0.53 y= 0+77.67/[1+(x/0.019)'2'34] 0.019
Macombl 0.74 y= 0+83.42[1+(x/0.027)'8-3°] 0.027

Monroe4 0.36 y= 0+60.99/[l+(x/0.041)‘l'82] 0.041
Oceanal 0.63 y= 0+82.16/[1+(x/0.11)'2'82] 0.11
Inghaml 0.44 y= 0+56.63/[1+(x/0.086)'25-""] 0.086
Macomb2 0.60 y= 0+63.33/[1+(x/0.11)"'38] 0.11
Monroe] 0.70 y= 0+61.36/[1+('x/0.12)‘3“3] 0.12
Newag02 b >0.138
Monroe2b >0.138

Bay] b >0.138

Ingham2 b >0.138
Masonl c

 

 

a All R2 values are signiﬁcant at 01 <0.01
b Population were highly tolerant and could not ﬁt logistic equation

° Population has huge variability in data which could not be transformed for analysis

61

Table 9. Log logistic equations, R2 values and GR50 values for eastern black nightshade

(SOLPT) populations in response to sulfosulfuron treatment.

 

 

SOLPT R 2: Log-logistic equation GR5n
Population (kg ai/ha)
Monroe2 0.83 y= 0+67.74[1+(x/0.032)'3'8] 0.032
Monroe] 0.73 y= 0+54.62[1+(x/0.038)'8'62] 0.038
Bay] 0.74 y= 0+66.95[1+(x/0.040)'8‘52] 0.04
Macomb2 0.73 y= 0+74.34[1+(x/0.053)"‘20] 0.053
Macombl 0.76 y= 0+91.24[1+(x/0.062)'0'75] 0.062
Monroe4 0.70 y= 0+80.99[1+(x/0.O66)‘0‘7'] 0.066
Inghaml 0.71 y= 0+83.81[1+(x/0.O74)"'12] 0.074
Newayg02 0.75 y= 0+92.84[1+(x/0.077)"'°5] 0.077
Masonl 0.79 y= 0+69.32[1+(x/0.07)"'23] 0.079
Ingham2 0.88 y= 0+85.70[1+(id0.074)"-""] 0.099
Macomb3 0.54 y= 0+52.64[1+(x/0.10)"-°7] 0.10
Oceanal b >0.138

 

 

a All R2 values are signiﬁcant at 01 < 0.01

b Population were highly tolerant and could not ﬁt logistic equation

62

Table 10. Log logistic equations, R2 values and GR50 values for eastern black nightshade

(SOLPT) populations in response to pyridate treatment.

 

 

 

SOLPT R 2“ Log-logistic equation GR50
Population (kg ai/ha)
Monroe2 b <0.22
Monroe4 b <0.22
Monroe] " <0.22

Newayg02 b <0.22

Macomb3 b <0.22
Macomb2 b <0.22
Macombl b <0.22

Masonl 0.60 y= 0+99.04[l+(x/0.29)'3'02] 0.29

Inghaml 0.70 y= 0+99.l6[1+(x/0.29)'2‘69] 0.29

Bay] 0.69 y= 0+100.00[l+(x/0.29)’3'45] 0.29
Oceanal 0.60 y= 0+98.59[1+(x/0.36)'2'32] 0.36
Ingham2 0.82 y= 0+100.00 [1+(x/0.44)'4‘75] 0.44

 

 

a All R2 values are signiﬁcant at a < 0.01

b Population were highly sensitive and could not ﬁt logistic equation

63

Table 11. Log logistic equations, R2 values and GRsn values for hairy nightshade

(SOLSA) populations in response to metribuzin treatment.

 

 

SOLSA R 2“ Log-logistic equation GRSO
Population (kg ai/ha)
Montcalm] 0.71 y= 0+ 82.17/[1+(x/0.10)’4‘89] 0.10
Montcalm2 0.85 y= 0+ 100.81/[1+(x/0.15)'2'06] 0.15

Presque2 0.90 =0+ 102.23/[1+(x/0.16)'2'73] 0.16
Presque3 0.88 y=0+ 96.85/[1+(x/0.19)“‘-8'] 0.19
Presque4 0.92 y=0+ 100.40/[1+(x/0.l9)'5‘33] 0.19
Presquel 0.91 y=0+ 99.97/[1+(x/0.20)'5-‘°] 0.20

Bay2 0.88 y=0+ 100.00/[]+(x/0.20)'2'87] 0.20
Macombl 0.86 y=0+ 95.47/[1+(x/0.23)'5’60] 0.23

 

 

a All R2 values are signiﬁcant at 01 < 0.01.

64

 

 

Table 12. Log logistic equations, R2 values and GR50 values for hairy nightshade

(SOLSA) populations in response to sulfentrazone treatment.

 

 

SOLSA R 2 Log-logistic equation GR50
Population (kg ai/ha)
Presque2 b <0.028
Presque3 b <0.028

Presquel 0.82 y= 0+ 100.59/[1+(x/0.04)‘°-77] 0.045
Montcalm2 0.83 y= 0+ 97.07/[1+(x/0.061)'2'22] 0.061
Montcalm] 0.77 y= 0+94.82/[1+(x/0.09)°‘05] 0.061

Presque4 0.86 y= 0+ 103.69/[1+(x/0.066)"-67] 0.066

Bay2 0.81 y= 0+ 82.56/[1+(x/0.070)"°-9'] 0.070
Macombl 0.69 3: 0+ 101.54/[1+(x/0.10)'0‘8l] 0.10

 

 

a All R2 values are signiﬁcant at 01 < 0.01.

b Population were highly sensitive and could not ﬁt logistic equation

65

 

Table 13. Log logistic equations, R2 values and GRso values for hairy nightshade

(SOLSA) populations in response to sulfosulfuron treatment.

 

 

SOLSA R 2“ Log-logistic equation GRso
Population (kg ai/ha)
Montcalm2 b <0.0086
Presquel b <0.0086
Presque2 0.60 y= 0+ 86.06/[1+(x/0.0099)'8-6‘] 0.0099
Presque3 0.81 y= 0+ 101.92/[1+(x/0.0099)"-39] 0.0099
Presque4 0.84 y= 0+ 97.84/[1+(x/0.0099)“‘-22] 0.0099
Bay2 0.91 y= 0+ 95.22/[1+(x/0.03 1)"'43] 0.031
Macomb] 0.87 y= 0+ 74.75/[1+(x/0.037)'2"72] 0.037
Montcalm] 0.84 y= 0+ 101.96/[1+(x/0.037)"-9‘] 0.037

 

 

a All R2 values are Signiﬁcant at a < 0.01.

b Population were highly sensitive and could not ﬁt logistic equation.

66

Table 14. Log logistic equations, R2 values and GRso values for hairy nightshade

(SOLSA) populations in response to halosulfuron treatment.

 

 

SOLSA R 2" Log-logistic equation GRSO
Population (kg ai/ha)

Presque2 0.59 y= 0+ 68.69/[1+(x/0.021)'2’48] 0.021

Presque3 0.28 y= 0+ 67.39/[1+(x/0.57)'°~37] 0.057
Montcalm2 0.70 y= 0+ 79.06/[1+(x/0.073)'0‘87] 0.073

Presque4 0.62 3: 0+ 79.58/[1+(x/0.077)'°-7"] 0.077

Bay2 b 0.31 >0.138

Macombl b 0.21 >0.138
Presquel °

Montcalm] °

 

a All R2 values are signiﬁcant at 01 < 0.01.

b Population were highly tolerant and could not ﬁt logistic equation

c Population has huge variability in data which could not be transformed for analysis

67

Table 15. Log logistic equations, R2 values and GRso values for hairy nightshade

(SOLSA) populations in response to pyridate treatment.

 

 

SOLSA R 2“ Log-logistic equation GR50
Population (kg ai/ha)
Montcalm] b <0.22
Montcam12 b <0.22
Presque2 b <0.22
Presque3 b <0.22
Presque4 b <0.22
Presquel b <0.22
Bay2 b <0.22
Macombl b <0.22

 

 

a All R2 values are Signiﬁcant at 01 < 0.01

b Population were highly sensitive and could not ﬁt logistic equation

68

V

 

Table 16. Log logistic equations, R2 values and GR50 values for horsenettle (SOLCA)

populations in response to halosulfuron treatment.

 

 

SOLCA R 2“ Log-logistic equation GR50
Population (kg ai/ha)
Monroe5 b <0.0086

Berrien2 0.83 y= 0+ 100.85/[1+(x/0.086)'2'5°] 0.0086
Montcalm3 0.60 y= 0+ 95.53/[1+(x/0.016)'°'38] 0.016
Vanburenl 0.42 y= 0+ 88.70/[1+(x/0.036)'°-47] 0.036

 

 

a All R2 values are signiﬁcant at 01 < 0.01.

b Population were highly sensitive and could not ﬁt logistic equation

69

Table 17. Log logistic equations, R2 values and GR50 values for horsenettle (SOLCA)

populations in response to sulfentrazone treatment.

 

 

SOLCA R 2“ Log-logistic equation GRso
Population (kg ai/ha)
Montcalm3 0.68 y= 0+ 80.81/[1+(x/0.012)"~54] 0.012

Berrien2 0.88 y= 0+ 70.70/[1+(x/0.12)"5"6] 0.12

Monroe5 0.85 y= 0+ 88.92/[1+(x/0.19)'2‘85] 0.19
Vanburenl 0.43 y= 0+ 65.07/[1+(x/0.35)“-‘°] 0.35

 

 

a All R2 values are signiﬁcant at a < 0.01.

70

Table 18. Log logistic equations, R2 values and GRso values for horsenettle (SOLCA)

populations in response to sulfosulfuron treatment.

 

 

SOLCA R 23 Log-logistic equation GR5n
Population (kg ai/ha)
Vanburenl b <0.0086
Montcalm3 0.68 y= 0+ 80.81/[1+(x/0.012)"-54] 0.012

Berrien2 0.49 y= 0+ 80.83/[1+(x/O.020)'°'30] 0.020

Monroe5 0.50 y= 0+ 88.91/[1+(x/0.031)'°'42] 0.031

 

 

a All R2 values are signiﬁcant at 01 < 0.01.

b Population were highly sensitive and could not ﬁt logistic equation

71

Table 19. Log logistic equations, R2 values and GR50 values for horsenettle (SOLCA)

populations in response tof pyridate treatment.

 

 

SOLCA R 2“ Log-logistic equation GR50
Population (kg ai/ha)
Monroe5 b <0.252

Berrien2 0.90 3: 0+ 103.29/[1+(x/0.40)"-7'] 0.40
Vanburenl 0.90 y= 0+ 93.3 8/[1+(x/0.40)'2'63] 0.40
Montcalm3 0.70 y= 0+ 98.44/[1+(x/0.51)"'0'] 0.51

 

 

a All R2 values are signiﬁcant at 01 < 0.01

b population were highly sensitive and could not ﬁt logistic equation.

72

Table 20. Log logistic equations, R2 values and GR50 values for horsenettle (SOLCA)

populations in response to metribuzin treatment.

 

 

SOLCA R 2“ Log-logistic equation GR50
Population (kg ai/ha)
Monroe5 b <0.070
Berriean <0.070

Vanburenl b <0.070
Montcalm3 b <0.070

 

 

a All R2 values are Signiﬁcant at a < 0.01

b Population were highly sensitive and could not ﬁt logistic equation

73

Table 21. Log logistic equations, R2 values and GR50 values for clammy groundcherry

(PHYHE) population in response to sulfosulfuron, metribuzin, pyridate, sulfentrazone

 

 

treatment.

PHYHE Treatments R 2“ Log-logistic equation GRSO
Population ( kg ai/ha)
Newagol Sulfosulfuron 0.54 y= 0+ 82.19/[1+(x/0.028)'°-42] 0.028

Metribuzin 0.73 y= 0+ 96.67/[1+(x/0.42)'0‘72] 0.42
Pyridate 0.62 y= 0+ 84.71/[1+(x/0.67)"'0°] 0.67
Sulfentrazone b >0.44
Halosulfuron c

 

 

a All R2 values are signiﬁcant at a < 0.01

b Population were tolerant to the treatment and could not ﬁt logistic equation

c Population has huge variability in data which could not be transformed for analysis

74

Table 22. Log logistic equations, R2 values8 and GRSO values for smooth groundcherry

(PHYSU) population in response to sulfosulfuro, pyridate, sulfentrazone treatment.

 

 

PHYSU Treatments R 2 Log-logistic equation GR50
Population (kg ai/ha)
Ingham3 Halosulfuron 0.58 y= 0+ 83.22/[1+(x/0.038)'1‘39] 0.038
Sulfosulfuron 0.61 y= 0+ 72.71/[1+(x/0.091)'°-"°] 0.091
Sulfentrazone 0.76 y= 0+ 102.21/[1+(x/0.104)'2"°] 0.104

Pyridate 0.69 y= 0+ 98.83/[1+(x/1.39)'0'70] 1.39
Metribuzin b <0.070

 

 

a All R2 values are signiﬁcant at a < 0.01

b Population were highly sensitive to the treatment and could not ﬁt logistic equation

75

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— — - Monroe4 (R2 = 0.78)

_______ Mason1 (R2 = 0.66)

Monroe1(R2= 0.82)
NewaygoZ (R2 = 053)

 

 

 

 

Figure 1a. Log logistic dose response curves of eastern black nightshade populations -

Monroe4, Masonl , Monroe], and Newag02 in response to metribuzin.

77

100

 

 

 

 

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0.0 0.2 0.4 0.6 0.8 1.0 1.2
Metribuzin (kg/ha)

 

— — — - Macomb2 (R2 = 0.61)
— -------- Macomb1 (R2 = 0.74)
.............. Macomb3 (R2 = 0,72)
—— Oceana1 (R2 = 0.047)

 

 

 

Figure 1b. Log logistic dose response curves of eastern black nightshade populations -

Macombl , Macomb2, Oceanal , and Macomb3 in response to metribuzin.

78

 

80

 

 

 

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Metribuzin (kg/ha)

 

one. 000000000 Oceana“ (R2 = 0047)
— — — Monroe 4 (R2= 0.78)

 

 

 

Figure 1c. Log logistic dose response curves Of the most susceptible - Monroe4 and

tolerant - Oceanal eastern black nightshade populations in response to metribuzin.

79

120

 

 

 

 

 

 

 

NewaygoZ (R2 = 0.36)
Macomb1 (R2 = 0.90)
Macomb2 (R2 = 0.76)
Macomb3 (R2 = 0.78)

 

 

 

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Sulfentrazone (kg/ha)

Figure 2a. Log logistic dose response curves of eastern black nightshade populations -

Newag02, Macomb3, Macombl, and Macomb2 in response to sulfentrazone.

80

100

 

     

 

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Sulfentrazone (kg/ha)

 

 

Monroe2 (R2 = 0.45)
Oceana1 (R2 = 0.75)
Mason1 (R2 = 0.91)

 

 

 

0.5

Figure 2b. Log logistic dose response curves of eastern black nightshade populations -

Oceanal , Monroe2 and Mason] in response to sulfentrazone.

81

120

 

   

 

 

 

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._ — — - Bay1 (R2 = 0.82)

.. ........ Monroe1 (R2= 0.79)

Ingham2(R2=0.67)
lngham1 (R2= 0.65)

 

 

 

Figure 2c. Log logistic dose response curves of eastern black nightshade populations -

Bay1, Monroe1, Ingham2, and Inghaml in response to sulfentrazone.

82

 

120

 

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Sulfentrazone (kg/ha)

 

Ingham2 (R2 = 0.67)
— Newaygoz (R2 = 0.36)

 

 

 

 

0.5

Figure 2d. Log logistic dose response curves Of the most susceptible Newag02 and

tolerant Ingham2 eastern black nightshade populations in response to sulfentrazone.

83

 

100

 

 

 

 

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0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 0.16
Halosulfuron (kg/ha)

 

— — — - Macomb3 (R2 = 0.53)
Macomb1(R2=0.74)
-- -------- Monroe4 (R2: 0.36)
------ Oceana1 (R2 = 0.63)
—— Macomb2 (R2 = 0.60)

 

 

 

Figure 3a. Log logistic dose response curves of eastern black nightshade populations -

Macomb3, Macombl , Macomb2, Oceanal , and Monroe2 in response to halosulfuron.

84

70

 

505

50

40

30

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20- ./'

Dry weight reduction (%)

10 - ./

 

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0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 0.16

 

Sulfentrazone (kg/ha)
_ _ —— - Monroe1 (R2 = 0.70)
------ lngham1 (R2 = 0.44)
ooeooocoooooo Neway902 (R2=018)
_ ........ Monroe2 (R2 = 0.63)

 

Bay1 (R2 = 0.49)

 

 

 

Figure 3b. Log logistic dose response curves of eastern black nightshade populations -

Monroe1 , Inghaml, Newag02, Monroe2, and Bay] in response to halosulfuron.

85

100

 

 

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Sulfentrazone (kg/ha)

 

NewaygoZ (R2 = 0.13)
— — — - Macomb3 (R2 = 0.53)
- -------- Macomb1 (R2 = 0.74)
.............. Macomb2 (R2 = 0.60)

 

 

 

Figure 3c. Log logistic dose response curves of the most susceptible Macomb3,
Macombl and tolerant Newag02, Macomb2 eastern black nightshade populations in

response to halosulfuron.

86

 

80

60-

40-

204

Dry weight reduction (%)

 

 

 

 

I I f I I

0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 0.16
Sulfosulfuron (kg/ha)

 

- — — Monroe2 (R2 = 0.83)

------ Monroe1 (R2 = 0.73)

............. Bay1 (R2 = Q74)
Macomb2 (R2 = 0.73)

 

 

 

Figure 4a. Log logistic dose response curves Of eastern black nightshade populations -

Monroe2, Monroe1 , Bay] and Macomb2 in response to sulfosulfuron.

87

100

 

80‘

60-

40*

Dry weight reduction (%)

 

 

 

 

I r I I I

0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 0.16
Sulfosulfuron (kg/ha)

 

— — — - Macomb1 (R2 = 0.76)
.............. Monroe4 (R2 = 070)
_ ........ Ingham1 (R2 = 0.71)
—— NewaygoZ (R2 = 075)

 

 

 

Figure 4b. Log logistic dose response curves of eastern black nightshade populations —

Macomb1, Monroe4, Inghaml and Newag02 in response to sulfosulfuron.

88

 

100

 

 

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0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 0.16
Sulfosulfuron (kg/ha)

 

_ _ — Mason1 (R2 = 0.79)

------- Ingham2 (R2 = 0.88)
—— Macomb3 (R2 = 0.54)
no... oooooooo Oceana1 (R2 = 047)

 

 

 

Figure 4c. Log logistic dose response curves Of eastern black nightshade populations -

Mason1, Ingham2, Macomb3, and Oceanal in response to sulfosulfuron.

89

80

 

 

60a

40-

20d

Dry weight reduction (%)

 

 

 

 

I I I I I I 1

0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 0.16

Sulfosulfuron (kg/ha)

 

 

Monroe2 (R2 = 0.83)
Oceana1 (R2: 0.47)

 

 

 

Figure 4d. Log logistic dose response curves of the most susceptible Monrer and

tolerant Oceanal eastern black nightshade populations in response to sulfosulfuron.

90

 

120

 

 

100 — ,:g;:a______

Dry weight reduction (%)

 

 

 

 

0 I I I I
0 1 2 3 4 5

Pyridate (kg/ha)

— — — - Mason1 (R2 = 0.60)

——————— Ingham1 (R2: 0.70)
.............. Bay1 (R2: 0.69)

_ ........ lngham2 (R2 = 0.82)
— Oceana1 (R2 = 0.36)

 

 

 

 

Figure 5a. Log logistic dose response curves of eastern black nightshade populations -

Mason1, Ingham2, Inghaml, Bay] and Oceana1 in response to pyridate.

91

120

 

 

 

 

 

 

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Metribuzin (kg/ha)

 

_. —— —- Montcalm1 (R2 = 0.71)
_______ Montcalm2 (R2 = 0.85)
0000000000000 Presquez (R2: 090)
—— Presque3 (R2 = 0.88)

 

 

 

Figure 5b. Log logistic dose response curves of hairy nightshade populations -

Montcalml , Montcalm2, Presque2, and Presque3 in response to metribuzin.

92

120

 

 

 

 

 

 

 

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— — — Presque4 (R2 = 0.92)
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—o-o-o- Bay2 (R2 = 0.88)
—— Macomb1 (R2 = 0.86)

 

 

 

FigureSc. Log logistic dose response curves of hairy nightshade populations - Presque4,

Presquel , Macomb1, and Bay2 in response to metribuzin.

93

120

 

 

 

 

 

 

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0.0 0.2 0.4 0.6 0.3 1.0 1.2

Metribuzin (kg/ha)

— Montca|m1 (R2 = 0.71)
Macomb1 (R2 = 0.86)

 

 

 

 

Figure 5d. Log logistic dose response curves of the most susceptible Montcalmland

tolerant Macombl hairy nightshade populations in response to metribuzin.

94

120

 

 

 

 

 

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Sulfentrazone (kg/ha)

 

_ —. — - Presque2 (R2 = 0.51)
— -------- Presque3 (R2 = 0.61)
—— Presque1 (R2 = 0.82)
Montcalm2(R2=0.83)

 

 

 

Figure 6a. Log logistic dose response curves of hairy nightshade populations - Presque2,

Presque3, Presque1 and Montcalm2 in response to sulfentrazone.

95

 

 

 

 

 

 

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0.0 0.1 0.2 0.3 0.4 0.5

Sulfentrazone (kg/ha)

 

— -— — - Montcalm1 (R2 = 0.77)
Presque4 (R2 = 0.86)
—.-—.-—-- Macomb1 (R2= 0.69)
Bay2 (R2 = 0.81)

 

 

 

 

Figure 6b. Log logistic dose response curves of hairy nightshade populations - Presque4,

Montcalm] , Macomb1, and Bay2 in response to sulfentrazone.

96

 

120

 

 

 

 

100 -
S‘:
c 80 -
.9.
‘5
3
"o
2 60 -*
E
.9
a)
3 40 q
2‘
Q

20 -*

O I— I I I
0.0 0.1 0.2 0.3 0.4 0.5

Sulfentrazone (kg/ha)

 

Presque2 (R2 = 0.51)
— — — Macomb1 (R2 = 069)

 

 

 

Figure 60. Log logistic dose response curves of the most susceptible Presque2 and

tolerant Macomb1 hairy nightshade populations in response to sulfentrazone.

97

110

 

 

 

 

 

100- ........................ .. ........
g 90-
C
.3 80-
o
3
13
2 70-
E
.9 60
m .—
3
2'
D 50—
40‘
30 u

 

I I I I I I
0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 0.16
Sulfosulfuron (kg/ha)

 

— — —- Presque2 (R2 = 0.60)
............. Presque3 (R2 = 081)
—-———— Presque4 (R2 = 0.84)

 

 

 

Figure 7a. Log logistic dose response curves of hairy nightshade populations - Presque2,

Presque3, and Presque4 in response to sulfosulfuron.

98

120

 

 

100‘ ooooooooooo 0 °°°°°
a", 80‘
I:

.2

‘3’ 604

U

9.’

5, 4o-

'5

3

E 20-
0+

 

 

 

0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 0.16
Sulfosulfuron (kg/ha)

 

Bay2 (R2 = 0.91)
_ — — Macomb1 (R2 = 0.87)
ooooooooooooo Montcalm“ (R2 = 084)

 

 

 

Figure 7b. Log logistic dose response curves of hairy nightshade populations - Macomb1,

Montcalm], and Bay2 in response to sulfosulfuron.

99

120

 

 

 

 

 

100-
g 80-
.5 / ————————————
8 so- I
'U
9 I
E» 40- I
(D
a I
E 20* I

/
0- __./
I I I I T I T

0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 0.16
Sulfosulfuron (kg/ha)

 

Presque4 (R2 = 0.34)
— — —- Macomb1 (R2= 0.87)

 

 

 

Figure 7c. Log logistic dose response curves of the most susceptible Presque4 and

tolerant Macomb1 hairy nightshade populations in response to sulfosulfuron.

100

 

100

30 4 -
O//

’3 ...e"
E, /,—-—-—'"—__';;Ty"’:.:::
'9 I’Tt‘).
..6 / ”’ .0 .
3 I, ..‘7
.0 / ’l’ ..gy .- —- __ — ‘—
9 40 - 9 /” O..‘./.’o". I'- .—
3 // ..’ ’3”
g If y'

20 - .z'
E‘ l /,/'
D /

O ..

 

 

 

 

I I I I j I I

0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 0.16
Halosulfuron (kg/ha)

 

_ _ — - Presque2 (R2 = 0.59)

------ Presque3 (R2 = 0.28)
n... ..... ..u Montcalm2 (R2 = 0.70)
_ ....... Presque4 (R2 = 0.62)
_ ........ Bay2 (R2 = 0.31)

 

 

 

Macomb1 (R2 = 0.21)

 

Figure 8a. Log logistic dose response curves of hairy nightshade populations - Presque2,

Presque3, Montcalm2, Presque4, Bay2. and Macombl in response to halosulfuron.

101

80

 

 

/”-— —————————
501 /
s /
.5 /
3401 l
e /
E /
a)
-- 20~
g /
Z‘
0
0—1

 

 

 

0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 0.16
Halosulfuron (kg/ha)

 

_ — — Presque2 (R2 = 0.59)
Macomb1 (R2 = 0.21)

 

 

 

Figure 8b. Log logistic dose response curves of the most susceptible Presque2 and

tolerant Macomb1 hairy nightshade populations in response to halosulfuron.

102

 

120

 

 

 

 

 

100 -
’o‘ —- -—’-""'
°\ ...— ‘__. ‘- -— ’°’
v ...— ’
c ’ I .’ ’ .....o
.9 ,z" °"'
'3 80 ‘ ,/" ..o'"
:r ./
'c
e .
3
2‘
D

40 -

20 I I I I I I I
0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14

Figure 9. Log logistic dose response curves of horsenettle populations - Berrien2,

Halosulfuron (kg/ha)

 

 

Berrien2 (R2 = 0.83)
Monroe5 (R2 = 0.50)
Montcalm3 (R2 = 0.60)
Vanburen1 (R2 = 0.42)

 

 

Vanburenl , Montcalm3, and Monroe5 in response to halosulfuron.

103

0.16

100

 

 

     

 

 

’ ’ .
80 —
I; oooooooooooooooo oﬂoooooooo 0000000000000000000
e: / ,
_ /"
,5 so / ./,.
g : / /°./
U : /'.
9 40 " : /
E : ,./'
.9 : .o/
G3) .° ./
20 - k:
b :9-
O .f'.’
,/' .'
l' / . .'
0 -1 “0000»...
0.0 0.1 0.2 0.3 0.4

Sulfentrazone (kg/ha)

 

 

Berrien2 (R2 = 0.88)
Monroe5 (R2 = 0.85)
Montcalm3 (R2 = 0.68)
Vanburen1 (R2 = 0.43)

 

 

0.5

Figure 10. Log logistic dose response curves of horsenettle populations - Berrien2,

Vanburenl , Montcalm3, and Monroe5 in response to sulfentrazone.

104

 

 

 

100
90- /
.‘E" 80-
CD
é
: 70~
.9
‘3
'o 60-
9
,5: 50-
G)
3
E‘ 40-
o
30~
20 I I I I I I I

 

 

 

0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 0.16
Sulfosulfuron (kg/ha)

 

oooooooooooooo Berrien2 (R2 : 049)
— — — Monroe5 (R2 = 0.50)
Montcalm3 (R2 = 0.68)

 

 

 

Figure 11. Log logistic dose response curves of horsenettle populations - Berrien2,

Montcalm3, and Monroe5 in response to sulfosulfuron.

105

120

 

 

 

 

 

 

 

100 -
is"
c 80 4
.9
‘6'
3
'0
93 60 -
E
.9
o
3 40 -
E“
D
20 a
o r . ' '
o 1 2 3 4 5
Pyridate (kg/ha)
Berrien2 (R2 = 0.90)
_. ........ MonroeS (R2 = 0.83)
_ _ — - Montcalm3 (R2 = 0.70)
00.00.000.000. Vanburen1 (R2: 090)

 

 

 

Figure 12. Log logistic dose response curves of horsenettle populations - Berrien2,

Vanburenl , Montcalm3, and Monroe5 in response to pyridate.

106

CHAPTER 4
EFFECT OF TEMPERATURE ON GERMINATION OF EASTERN BLACK
NIGHTSHADE, HAIRY NIGHTSHADE, HORSENETTLE, SMOOTH
GROUNDCHERRY AND CLAMMY GROUNDCHERRY

Abstract. Populations of eastern black nightshade (Inghaml, Oceana1), horsenettle
(Oceana4, Vanburenl), hairy nightshade (Bay1, Presque2), smooth groundcherry
(Ingham3) and clammy groundcherry (Newagol) were subjected to four different
day/night temperature regimes (28/20 °C, 24/ 16 °C, 22/ 14 °C, 15/10 0C) in the growth
chamber for 15 days with 14 hr day and 10 hr night at constant light intensity to analyze
the effect of temperature on germination of nightshade and groundcherry species.
Temperature regime had a signiﬁcant effect on germination of eastern black nightshade,
horsenettle, smooth groundcherry, and clammy groundcherry. Under these conditions
hairy nightshade populations did not germinate under any temperature regime. Eastern
black nightshade populations germinated at least 70% at 28/20 °C. Horsenettle
populations germinated at least 30% at 28/20 °C. Smooth groundcherry and clammy
groundcherry had germination of 29% and 8% respectively at 28/20 °C. Within eastern
black nightshade populations, Oceana1 germinated at a wider range of temperatures than
the Inghaml population.

Key words: Eastern black nightshade (Solanum ptycanthum), hairy nightshade (Solanum
sarrachoides), horsenettle (Solanum carolinense), smooth groundcherry (Physalis

subglabrata), clammy groundcherry (Physalis heterophylla), temperature.

107

INTRODUCTION

Nightshades are annual and short lived perennials that are common weeds in many parts
of the world. In the US. there are 11 recognized species (Schilling, 1981) with 4 species
in the Solanum nigrum complex recognized as troublesome weeds in Solanaceous crops
such as tomato, potato and pepper (Heiser, 1979). These species are eastern black
nightshade (Solanum ptycanthum), black nightshade (Solanum nigrum), American black
nightshade (Solanum americanum), and hairy nightshade (Solanum sarrachoides) (Ogg et
al., 1981). In the U.S., eastern black nightshade is found primarily east of the Rocky
Mountains (Bassett et al., 1984), hairy nightshade is distributed throughout the U.S., and
black nightshade and American black nightshade are found primarily in the Western
(Ogg et al., 1981) and southern US. In addition to these annual nightshades, perennial
nightshades such as horsenettle (Solanum carolinense) occur throughout the eastern US.
Groundcherries such as clammy groundcherry (Physalis heterophylla) and smooth
groundcherry (Physalis subglabrata) also are found throughout the eastern US.

In the north eastern U.S., Solanaceous vegetable crop production (tomato, pepper,
eggplant, potato) is an important enterprise for many farmers. However yield loss due to
weeds has always been a challenge to farmers and weed scientists. Bridges (1992)
estimated about a 6.4 million dollar yield loss per year in fresh market tomato due to
weeds in Michigan. Eastern black nightshade, hairy nightshade, and horsenettle are major
competitive weeds in tomato production. Groundcherries such as smooth groundcherry
and clammy groundcherry, usually are not major competitive weeds compared to other

nightshade species.

108

The shift in the distribution and abundance of nightshade species (in particular
eastern black nightshade) in the north eastern US. poses a continuing challenge to those
concerned with weed control, as well as to those seeking to understand the causes of
these shifts. Evolutionary changes have played a key part in the changing distribution
pattern of nightshade species (Hermanutz and Weaver, 1990). Germination requirements
of nightshade species is an important factor in species dominance and adaptation to
dynamic environments.

Knowledge of germination patterns of different nightshade and groundcherry
species is important in planning effective weed control programs. There is a typical
period (or periods) of high emergence that is characteristic for each weed species
(Brenchley and Warington, 1930; Chepil, 1946; Roberts, 1964; Roberts and Feast, 1970;
Stoller and Wax., 1973). The time of weed seedling emergence can, in part, explain
which species will be the most serious weeds within a given crop management system.
Stoller and Wax (1973) concluded that weeds that complete most of their emergence
early are likely to be killed during soil preparation before planting corn or soybean and
are not seriously troublesome weeds after corn planting.

There are many germination studies determining the period of emergence of
eastern black nightshade in US. Ogg and Dawson (1984) reported that eastern black
nightshade seedlings began emerging in Washington in late March, with most emerging
in April. Quakenbush and Andersen (1984) reported that eastern black nightshade
germination began in mid April and early May in Minnesota. Studies by Ogg and
Dawson (1984) reported that hairy nightshade germinated in Washington in late March to

early April of each year and continued to emerge throughout the growing season. In

109

Warwick (U.K.), hairy nightshade germination ranged from April to June and induced
dormancy occurred during August (Roberts and Boddrell, 1983). In general, soil
temperatures optimum for germination of eastern black nightshade, hairy nightshade and
smooth groundcherry are between 25-30°C (Vandeventer et al., 1982; Roberts and
Boddrell, 1983; Thomson and Witt, 1987). But there was no information reported on the
optimum temperature for germination of clammy groundcherry and horsenettle.

Because of the prevalence and importance of nightshade and groundcherry
species in Michigan and the need for biological information to establish effective
management systems, studies were conducted to obtain information on the inﬂuence of
temperature on germination of eastern black nightshade, hairy nightshade, horsenettle,

smooth groundcherry and clammy groundcherry.

110

MATERIALS AND METHODS:

Growth chamber experiments were conducted twice during April 2003 to
determine the inﬂuence of temperature on the germination of nightshade and
groundcherry species. Two populations from each nightshade species and one population
from each groundcherry species were tested in this study. The nightshade populations
were selected on the basis of unique morphological features observed during collection of
nightshade berries in Michigan from different counties. The list of the population,
location and associated crop are presented in Table 2.

Processing of seeds:

The nightshade and groundcherry berries were taken to the laboratory at Michigan
State University and were crushed and soaked in water overnight to allow the fruit juices
to ferment. The seeds were then placed in cheese cloth and squeezed to remove as much
juice as possible. Then the seeds in the cheese cloth were washed repeatedly and air dried
for a week at normal room temperature (24 oC). The seeds of all the nightshade and
groundcherry populations were stored for two months at -20°C and then stored in an
incubator at 28° C to overcome dormancy.

Viability test:

A viability test was conducted a week before the ﬁrst germination study. Twenty
ﬁve seeds of each population were subjected to a tetrazolium (0.1%) test to determine the
viability of the seeds. The cotyledons of the seeds were cut into two halves (without
injuring the embryo) to expose the seeds to tetrazolium solution. The seeds were soaked
in 0.1% tetrazolium chloride solution in a Petri dish for about 3 hours. Seeds with

embryos that turned pink in color were considered to be viable and seeds with embryos

111

that did not change color were considered to be non-viable. The viability test results are
presented in Table 2.
Design of the experiment:

The experiment was designed as a randomized complete block with 4 replications
per treatment. Each replicate contained 50 seeds in one Petri dish. The base of the Petri
dish was covered with Whatmann No.50 filter paper to hold the moisture in the Petri dish
and to provide a base to anchor the seeds. Approximately 5ml of distilled water was
added to the Petridish every day as needed to moisten the seeds. The seeds were covered
by a thin, soft and porous white tissue paper to protect the seeds from the splashing effect
of water.

Treatments:

Four different day/night temperature regimes, 28/200 C, 24/ 16 ° C, 22/14 0 C and
15/10 0 C were set up in different growth chambers1 with a 14 h day and 10 h night. The
temperature and duration of day/night was selected in accordance with Michigan climatic
conditions during the summer. Light intensity of 35 uE/mz/sec and quality of 0.7 Red/Far
red ratios were maintained in all the growth chambers. The humidity in the growth
chamber was maintained at 80%.

Germination count:

The germination experiments were maintained for 15 days. The germination
counts were taken every 3 days. Seeds that had both a radicle and plumule were
considered germinated. The germinated seeds were removed from the Petri dishes after

counting and were discarded.

 

lConviron E15, Controlled environments Ltd.. Winnipeg. Manitoba, Canada.

112

Statistical anlaysis:

The data were subjected to analysis of variance (ANOVA) and germination
means of the nightshade and groundcherry species at the speciﬁc temperature regime
were separated by using Fisher’s protected LSD at a = 0.05 by using SAS program

(SAS, 1990).

113

RESULTS AND DISCUSSION:
Germination response of eastern black nightshade populations:

Signiﬁcant variation in germination percent was observed among eastern black
nightshade populations across temperature regimes except at 28/20 0C (Table 1).
Inghaml and Oceana1 populations had mean germination percent of 80 and 74 at 28/20
0C. However a steep decline in germination percent to 23, 7, and 0 % was observed in
Inghaml population at 24/16 °C, 22/ 14 °C, 15/10 0C respectively.

On the other hand, Oceana1 population had a mean germination percent of 74, 60,
24 at 24/20 °C, 22/ 14 °C, and 15/10 °C respectively. Clearly, Oceana1 population has a
potential of germination at wider range of temperature than Inghaml population and can
have a signiﬁcant effect on weediness during the growing season. Having the potential to
germinate early in the summer season at a temperature of 15/10 oC, Oceana1 eastern
black nightshade has the potential to increase vegetative growth due to lack of
competition from the crop and other weeds, during the early growing season. And hence
Oceana1 eastern black nightshade can reproduce early in the growing season and can
increase the seed bank in the soil within a short period.

There was considerable difference in period of emergence among eastern black
nightshade populations. Early germination was observed in the Oceanal population at
28/20 0C, 24/ 16 °C, 22/ 14 °C at day 3 and the maximum germination percent during the
germination count period was observed at day 6 across all temperature regimes (Figure
1). Due to rapid germination potential, the Oceana1 population can dominate other weeds
and compete strongly with the main crop for nutrients, light, space and water during the

growing season.

114

On the other hand, germination of the Inghaml population was observed 6 days
after seeding (DAS) in the Petri dish across all temperature regimes except at 15/ 10 °C.
Inghaml population had maximum germination percent at 9 DAS in the Petri dish
(Figure 2). Zero percent germination was observed in the Inghaml population during the
entire counting period at 15/ 10 °C. The Oceana1 population germinates earlier than the
Inghaml population.

Hermanutz and Weaver, (1990) observed similar result in northern populations
(Harrow, Wright, Pelee, and Rondeau) of Canada possessing broader range of
germination at various temperature regimes. They suggested that the colonization
potential of northwardly migrating eastern black nightshade population might be due to
genetic variability in germination parameters between northern and southern eastern
black nightshade populations. Hence genetic variability in germination parameters
between Inghaml and Oceana1 population may be one of the reasons for differences in
pattern across temperature regimes. From the results, the Oceana1 population has the
colonization potential in new habitats of Michigan due to early germinating characteristic

at wider range of temperature.

Germination response of horsenettle populations:

Horsenettle populations had no variation in germination across temperatures.
Oceana4 and Vanburenl populations had mean germination percentage of 37 and 33 at
28/20 0C respectively (Table 1). Very low germination is noted at other temperature
regimes. Previous reports on the inﬂuence of temperature on germination of horsenettle

seeds suggest that decreasing the day temperature and increasing the night temperature to

115

about 20/30 0C and 20/35 °C for 16/8 hours increased the germination of horsenettle to
84% and 87% respectively (Buhler and Hoffman, 1999).

Both the horsenettle populations were slow to germinate and there was no
signiﬁcant variation in time of germination among the populations. Oceana4 and
Vanburenl populations had maximum germination percent at 15 DAS at 28/20 0C
(Figure 4, 5). Based on the late and poor germination characteristic, horsenettle may be
less competitive and may not be as serious a weed as the other annual Solanaceous weeds

if horsenettle reproduces only through seeds.

Germination response of hairy nightshade populations:

None of the hairy nightshade populations germinated at any temperature (Tablel).
Induced dormancy might be the reason for zero percent germination. The following
description about induced dormancy was based on Booth et al.(2003).

Induced dormancy or secondary dormancy is developed by the dispersed seed in
response to the environment. Induced dormancy is usually imposed when environmental
conditions are unfavorable for prolonged periods of time. Induced dormancy is also
called a cyclical dormancy or seasonal dormancy, where seeds may cycle in and out of
dormancy, changing from dormant to conditionally dormant (where the seeds germinate
under a smaller range of conditions) to non-dormant; this cycle repeats and can result in
seasonal dormancy cycles. Seasonal dormancy is induced by temperature.

Many reports suggest that induced dormancy is one of the germination
characteristic of hairy nightshade. Roberts and Boddrell, (1983) reported that seasonal

dormancy of hairy nightshade lasted until late April or early May. Germination virtually

116

ceases at the end of August and develops back to induced dormancy at Warwick (U.K.).
Studies by Ogg and Dawson (1984) observed the seasonal dormancy characteristic of
hairy nightshade in Washington until late March or early April. The seasonal emergence
pattern of hairy nightshade, can maintain a persistent seed bank in the soil. During the
ﬁrst year, 30-45% of hairy nightshade seed in the seed bank can germinate and the others

continued to appear for at least next 4 years (Roberts and Boddrell, 1983).

Germination response of smooth groundcherry and clammy groundcherry:

Low germination percent was observed across temperature regimes for both
groundcherry species (Table 1). However, smooth groundcherry had a signiﬁcantly
higher germination percent than clammy groundcherry at 28/20 0C (Table 1). A gradual
decrease in germination across temperature regimes was observed in smooth
grouncherry. Both species are slow to germinate. Smooth groundcherry had maximum
germination at 15 DAS across all temperature regimes except at 15/ 10 °C (Figure 3).
Maximum germination of 5% was observed after 12 DAS at 28/20 °C (Figure 3). Poor
and slow germination of groundcherry species may cause less weediness than annual
nightshade species like eastern black nightshade and hairy nightshade if groundcherry
species reproduces only through seeds. In addition cultural practices followed in
vegetable crop production like tillage, herbicide application affects the germinated
groundcherry species produced from seeds.

The invasion success of a colonizing weed depends on the ability of its seed to
germinate in new habitats (Groves, 1986). Temperature regime constitutes one of the

most potent selective forces in limiting the spread of weeds in new habitats (Thompson,

117

1970). Based on these results, eastern black nightshade germinates earlier and at a higher
germination percentage than other nightshade and groundcherry species across
temperature regimes. Variability in germination characteristic of eastern black nightshade
populations has tremendous impact on future range of expansion and it is more likely to
be the dominant weed in the agriculture system. The seasonal emergence pattern of hairy
nightshade can lead to persistent problem in the ﬁeld for many years and it may become a
more common weed in agriculture. A crop rotation system and proper application timing

of pre emergence herbicides can be the better management system to control nightshades.

118

10.

Literature cited

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ptycanthum Dun., Snigrum L. and S.sarrachoides Sendt. Canadian Journal of
Plant Science 65: 401-414.

Booth, B.D., S.D. Murphy and C]. Swanton. 2003. Weed ecology in natural and
agricultural systems. CABI publishing, Cambridge, MA pp.92

. Brenchley, W.E. and K. Warington. 1930. The weed seed population of arable

soil. 1. Numerical estimation of viable seeds and observations on their natural
dormancy. J. Ecology 18: 235-272.

Bridges. 1992. Crop losses due to weeds in the United States. Allen Press,
Lawrence, Kansas pp. 30

Buhler, DD, and ML. Hofﬁnan. 1999. Andersen’s guide to practical methods of
propagating weeds and other plants. Allen Press, Lawrence, Ks pp. 128.

Chepil, W.S. 1946. Germination of weed seeds. II. The inﬂuence of tillage
treatments on germination. Science of Agriculture 26: 347-3 57.

Groves, RH. 1986. Invasion of Mediterranean ecosystems by weeds. In:
Resilience in Mediterranean- type ecosystems. Edited by B.Dell, A.J.M.Hopkins
and B.B.Lamont. Junk Publishers, Boston, MA pp.129-145.

Heiser, C.B., J., D.L. Burton and BE. Schilling, Jr. 1979. Biosystematics and
taxonomic studies of the Solanum nigrum complex in eastern North America.
The biology and taxonomy of the Solanaceae. Academic Press, NY pg 513-527.

Hermanutz, LA. and SE. Weaver. 1990. Germination and growth of Solanum
ptycam‘hum and Solnaum sarrachoides. Canadian Journal of Plant Science
71:167-174.

Lamondia, J.A., Smith, V.L., and SM. Douglass. 1999. Host range of Oidium
lycopersicum on selected Solanaceous species in Connecticut. Plant Disease
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119

 

m; -.=-..‘ ' tin-Eel...

11.

12.

13.

14.

15.

16.

17.

l8.

19.

20.

McGiffen, Jr., M.E., J.B. Masiunas and J. Hesketh. 1992. Competition for light
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Ogg A.G., Jr., B.S. Rogers and BE. Schilling. 1981. Characterization of black
nightshade (Solanum nigrum) and related species in the United States. Weed
Science 29:27-32.

Ogg, A.G., Jr. and J .H. Dawson. 1984. Time of emergence of eight weed species.
Weed Science 32: 327-335.

Perez, F.G.M. and J.B. Masiunas. 1990. Eastern black nightshade (Solanum
ptycanthum) interference in processing tomato (Lycopersicon esculentum). Weed
Science 38:385-388.

Quackenbush LS. and RN. Andersen, 1984. Distribution and biology of two
nightshades (Solanum spp.) in Minnesota. Weed Science 32: 529-533.

Roberts, HA. 1964. Emergence and longevity in cultivated soil of seeds of some
annual weeds. Weed Research 4: 296-307.

Roberts, HA and J.E. Boddrell. 1983. Field emergence and temperature

requirements for germination in Solanum sarrachoides. Weed Research 23: 247-
252.

Roberts, H.A. and RM. Feast. 1970. Seasonal distribution of emergence in some
annual weeds. Exp. Horticulture 21: 36-41.

[SAS] Statistical Analysis System. 1990. SAS Procedures Guides. Version 6, 3rd
edition, Cary, NC: Statistical Analysis Systems Institute 567 p.

Schilling, BE. 1981. Systematics of Solanum sect. Solanum (Solanaceae) in
North America Systematic Botany 6: 1 72-185.

120

21.

22.

23.

24.

Stoller, E.W. and L.M. Wax. 1973. Periodicity of germination and emergence of
some annual weeds. Weed Science 21: 574-269.

Thompson, PA. 1970. Characterization of the germination response to
temperature of species and ecotypes. Nature (London) 255: 827-831.

Thomson, EC. and W. Witt. 1987. Germination of cutleaf groundcherry (Physalis
angulata), smooth groundcherry (Physalis virginiana) and eastern black
nightshade (Solanum ptycanthum). Weed Science 35:58-62.

Vandeventer, J.W., W.F. Meggitt, and D.Penner.l982. Morphological and

physiological variability in black nightshade (Solanum spp) Pesticide Science
13:257- 262.

121

Tablel. Comparison of germination mean percent across temperature to analyze the trend
in variability among eastern black nightshade (SOLPT) populations, horsenettle
(SOLCA) populations, hairy nightshade (SOLSA) population, smooth groundcherry

(PHYSU) and clammy groundcherry (PHYHE) species.

 

Species

SOLPT
SOLPT
SOLCA
SOLCA
SOLSA
SOLSA
PHYHE
PHYSU

Population

In gham 1
Oceana1
Oceana4
Vanburen 1
Bay]
Presque2
Newagol

Ingham3

Standard deviation

LSD

Germination mean (percent)

 

28/20
(°C)

80a
74a
37b
33b
CC
CC
8c
29b

29.22
10.16

24/16

(’0

23b
74a
2d
1d
0d
0d
1d
1 0c

24.68

7.21

22/14
(’0

7b

60a

0b
4b
Ob
0b
0b
3b

20.11
6.85

15/10

(’0

0b
24a
0b
0b
0b
0b
0b
0b

7.96
5.77

 

Means followed by the same letter in the same column are not statistically signiﬁcant at a

= 0.5

122

Table2. List of eastern black nightshade, hairy nightshade, horsenettle, clammy
groundcherry and smooth groundcherry populations collected from various regions in
Michigan; including location, associated crap and percent seed viability.

 

 

Species Populations Regions collected from Current Viability
Michigan crop '
Eastern black Inghaml T3N, RIW, Sec212, Tomato 92%
nightshade Holt twp
Oceana1 T15N, R17W, Secz32, Asparagus 88%
Hart twp
Hairy Bay1 T15N, R3E, Secz22, Potato 92%
nightshade Garﬁeld twp
Presque Isle 2 T34N, R6E, Sec:30, Dry bean 88%
S.Pulawski twp
Horsenettle Vanburenl T3S, R16W, Sec: 1 , Tomato 84%
Hartford twp
Oceana4 T16N, R17W, Secz2, A* 92%
Weare twp
Clammy Newaygol T12N, R14W, Sec236, Tomato 88%
groundcherry Sheridan twp
Smooth Ingham3 T2N, R2W, Sec: 14, Onion 84%
groundcherry Aurelius twp

 

 

A*- Population is collected from uncultivated site.

123

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

so i; 1 so 1
= ‘5 1 z ‘5 1
7?: 4o; ‘ .3 4o
5 35 E 35 l
E230 . ‘ $230 1
2' § 25 " 1 Z§25
3.: 20 ‘- 3020 l
5. 1 a
1; 15 ., 1 f 15 l
c 1 II
8 5 ‘ = 5 " . .22 1
o l o o .1 - .- ¢¢‘- u,__ ... I
1 3 6 9 12 15
P071011 01 ”MIMI“ (DIV!) It W20 9 l Period of germination (Days) at 24116 C l.
(A) (B)
50 1
= 45 50
.3 .1 g 1.
E 35 I; 40
E .E
{:30 15,35
Go :“30
3:20 5 E
E" «1‘20
.; 15 E 15
5 1° E 10 l
II II
3 5 s 5 1
0 o o
6 9 12 15 3 6 9 12 15
1’9"“ 0‘ 90mm” 1°11") '1 ”I“ c Period of germination (Days) It 15110 c
1_ g , , ,_ Q
(C) (D)

Figure 1.Temperature effect on the period of germination of Oceana1 eastern black
nightshade population at A) 28/20° C, B) 25/ 16° C, C) 22/ 14° C and D) 15/ 10° C

124

 

 

 

 

 

 

 

 

 

 

 

 

 

50 50
45 , i , L 2 E 45 - - - 2
1: 3
g 40 L, ...... - E 40
g 35 g 35 , ,
E: 30 § 30 9 2 -
a: .E
c 2 25 E 25 i v
z o _ a
n. 20 ca 20 2 i
E" c
" 15 g 15
E E
5 5 E 5
0 E 0
3 6 9 12 15
Period 0f 90"“ "“110“ (ﬂy!) it 23120 c 1 Period of germ Inatlon (Days) at 2411sc
.-_l 1
(A) (B)
50 ---------
‘3 45 , - u - -
§
40 ,
a
g 35 2
§ 30 -
.5 25
E ,2 A
11'» 20 ~
1:
E 15 , e
.. 1o - W"
E
2 V. V V V
g 177-: {.91 7771
3 6 9 12 15

Period of germination (Days) at 22I14C

 

(C)

Figure 2. Temperature effect on the period of germination of Inghaml eastern black
nightshade population at A) 28/20 °C, B) 25/ 16 °C and C) 22/14 °C

125

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

50 — 50 —
1 c 45 a- g a», ,5 45 _ - g
'2-2 40”“. a _ - g 40~——— ————

i .5 35 _ A- u _. *m E 35 ~ , , v —
1§§301--_ ——— __._,W a; gE30«———— w A-— ~— ~
a 25 a- _____ _A-m. __.-__~ ‘ c 8 25 — — — v H

c § .. -.
8 a 20 ,___M ,,_ 0: 20 r- ﬂ m L ,
5.. 2-15 ____., _ _ﬂ_ﬂ mg
‘ E 1 5 P— >‘ w Am -—V H——.- 'w-i “Vi “A. 3 1 0 _ ‘— _.__. “— .._. ! __
. 2 10 m - ____ _ ._. m _- g 5
i ’— —__n *‘ PPPPPP A... _— —- g —— ,____ _ - ._A .. .. .— , r
3 6 9 12 15 3 5 9 12 15
Period of germination (Days) at 24(16 C Period 0' germination (Days) at 23,20 C
(A) (B)
50 w j .. 5° ‘ ' ‘ ‘
c 45 W _ _- - ____ _ _‘_ E 45 -, — - m— — ~—— w—«-
~+ -—- —- w— ~+ -—~—.
5 35 ___ —- — —~- # -_— ‘g 35 b _ ~ — H ~ m;
gawk “A .g _ €30» __ ._ _a m
as .E
5 e 25 — — F ..... _ _.— __ E 25 a _ W
Eézo-* W A,é - 3201 w , M-,
n15-—>-— —— E15 r _ - ”-4
25 1o - —- _——-———— 5 10 H __ ________ g.
"' r
m [771 l a y‘
1 5 m a-r: 1
3 6 9 12 15 3 6 9 12 15
Period of germination (DIV!) at 221146 Period or germ Inatlon (Days) at 24mm
(C) (D)
_ — * ﬂ 1
A 50 H V — m_ A
E, 45 M g *9 a
U
1 a 40 ..... - ___ _ __
i :35 m 7 A
O
. 7.; 30 «A 7 _
.E
E 25 4- 7 w a?“
1 8.20%» _ ____.
i 5 15 _
I E10 if. ____
w n
: E 5 L...” -w- _~_-..,ﬂ
_ 6 9 12 15

Figure 3. Temperature

( 3

Period of germination (Days) at 28I20 C

 

(E)

effect on the period of germination of clammy groundcherry
population Newagol at A) 24/16 0C, B) 28/20 OC and smooth groundcherry population

Ingham3 at C) 22/14 °C, D) 24/16 0C, B) 28/20 0C.

126

 

 

 

 

 

 

 

 

 

 

 

 

so 7 7777 777777777 77 50 .1
g 45 7 77 7 7 .3 45 7 7 77 7
=3 4 7 77 7 77 7 ..... 7 r” .- d C , V;
g 0 g 40 3
E 35 7 7 7 7 7 E 35 7 77 7 77 7 77777 7;
a?” 77 77 7 7 7 a? 30 7 7
§225 77 777 7 E22557 7 7 777:
5&2077 77 7 77 7 7 5&207 — — 77‘
g 15 _. __ .._. --._- _ __M a. '- 15 7 7 7 77
2 E 10 _ t ”a k _ _
a 10 —- 77 77777 a :3 77
a a :
c _ A_‘ 5 A F , _ a
0 ’— “‘ g ;7
> 7 r : > 0 .._... 7;
a 6 9 12 15 7 3 6 9 12 15
Period of germination (Days) at 28120 c Period of germination (Days) at 24116 C

 

(A) (B)

 

 

‘ ,_ ____ ,a _i _ __ A

E 45 . #_,a .. ____4

1 ‘a’ 40 - 77 77 ~ 7 7777- 7- 7 7;

.5 35 r, ,7 ,__~. . 7. .- v7 ._,..____ a."

E 3 30 1.-.. .._-_.__. .._ __ _____.._.. .___._.i

a 5 ;

g 2 25 7777777777777 ~ 7 ~- 777-- 7—7—-- - 777

g: 20 777-k -7777 A——-- 77:

l .- " i

‘ S 15 _ .__.-. . _ .--......_-_. ...__.~__

1 a 10 7—----7-— 777 - 7 7 77--—7-

: g 5 .. 77 f f- 7
‘ 3 6 9 12 15

Period of 9a rm Ination (Days) at 22I15 C

 

 

(C)

Figure 4. Temperature effect on the period of germination of Vanburenl horsenettle
population at A) 28/20 °C, B) 25/ 16 °C, and C) 22/14 0C

127

 

 

 

 

 

 

 

 

 

5 S 45 ' ’ ’ ’ * i ’ i ’
E '7; 4o 7 7 —
C
E ‘g 357777 77777777
8.55? ggso 7 77 7 7 7 7
5 2 § §’ 25 7 7 7 7 7
3g Egzo 7 7 7 7 7 7 7
8 8 10 7 77 7 777 77 7
O 0
O o 5 7 7 2- ﬂ 7. W, H A
0 urn I!“
3 6 9 12 15 ‘ 3 6 9 12 15 ‘
Period of gemination (Days) at 28/200 1‘ Period of germ‘nation (Days) at 24/160 1
i
,.v _ A1 --_,,- ___ _4
(A) (B)
if a, , ,, , , i _
i 50 -- - — 77 7 7— 7
‘ S 45 777777 77 7 7 7 ~7
'75 40 7 7 7 7 7 7
‘ g 35 7 7 7 7 77 77 7
‘.=307 7 77 77777 77
a:
i 5 § 25 777 7 7 7 777 7 7 7
J E5": 20 77 777 ~ 7 7 77 7 7
¢ 15 7 7 7 7 7
I
i 10 777 7 7 7 7 7 777
o 5 7 7 77 77 777 777 77
§ 3 6 9 12 15
\ Period of germination (Days) at 22J14 C

 

 

(C)

Figure 5. Temperature effect on the period of germination of Oceana4 horsenettle
population at A) 28/20 °C, B) 25/16 °C, and C) 22/ 14 °C

128

APPENDICES

List of common and chemical names of herbicides (Weed Science Society of

America, 2002)

 

Common name

Trade name

Chemical name

 

Aciﬂuorfen

Carfentrazone

DCPA

Dimethenamid- P

Diquat

Flumioxazin

Halosulfuron

Imazethapyr

Imazamox

Methyl bromide

Metribuzin

Blazer

Aim

Dacthal

Outlook

Reglone

Valor

Sandea

Pursuit

Raptor

Methyl
bromide,
Pic Brom 25
Sencor

5-[2-chloro-4-(triﬂuoromethyl)phenoxy]-2-
nitrobenzoic acid

X,2-dichloro-5-[4-(diﬂuromethyl)-4,5-
dihydro-3 -methyl-5-oxo-lH-1 ,2,4-triazol-1 -
yl ]-4-ﬂuorobenzenepropanic acid

Dimethyl 2,3,5,6-tetrachloro-1 ,4-
benzenedicarboxylate), and chloramben (3-
amino-2,5-dichlorobenzoic acid

2-chloro-N(2,4-dimethyl-3-thienyl)-N-(2-
methoxy-l -methylethyl)acetamide

6,7-dihydrodipyrido[ 1 ,2-a22’ ,1 ’-
c]pyrazinediium ion

2-[7-ﬂuoro-3,4-dihydro-3-oxo-4-(2-propnyl)—
2H-1 ,4-benzoxazin-6yl] -4,5 ,6,7-tetrahydro-
1H-isoindole-l,3(2H)-dione

3-chloro-5[[[[(4,6-dimethoxy-2-
pyrimidinyl)amino]carbonyl]amino]sulfonyl]-
1-methyl-1H-pyrazole-4-carboxylic acid

2- [4,5-dihydro-4-methyl-4-( 1 -methylethyl)—5 -
oxo-lH—imidazol-Z-yl]-5-ethyl-3-
pyridinecarboxylic acid

2-[4,5-dihydro-4-methyl-4-(1 -methylethyl)-5 -
oxo-1H—imidazol-Z-yl]-5-(methoxymethyl)-3-
pyridinecarboxylic acid

bromomethane

4-amino-6-(1 ,1 -dimethylethyl)-3- ‘
(methylthio)-1 ,2,4-triazin-5 (4H)-one

 

129

 

Common name

Metolachlor-S Dual II
magnum
Napropramide Devrinol
Oxyﬂourfen Goal
Paraquat Gramoxone
Primisulfuron-methyl Beacon
Pyridate Tough
Rimsulfuron Matrix
Sulfentrazone Spartan
Sulfosulfuron Maverick

Chemical name

2-chloro-N-(2-ethyl-6-methylphenyl)-5,7-
dimethoxy[1,2,4]triazolo[l,5-a]pyrimidine-2-
sulfonamide

N, N-diethyl-2-( 1 -naphthalenylox y)
propanamide

2-chloro-1 -(3-ethoxy—4-nitrophenoxy)-4-
(triﬂuoromethyl)benzene

1,1 ’-dimethyl-4,4’-bipyridinium ion

2-[[[[[4,6,-bis(diﬂuoromethoxy)- l ,3 ,5-
triazine-2-yl]amino]
carbonyl]amino]sulfonyl]benzoic acid

0-(6-(chloro-3 -phenyl-4-pyridazinyl) S—octyl
carbonothioate

N-[[(4,6-dimethoxy-2-
pyrimidinyl)amino]carbonyl]-3-
(ethylsulfonyl)-2-pyridinesulfonamide

N-[2,4-dichloro-5-[4-(dichloro-5-[4-
(diﬂuromethyl)-4,S-dihydro-3-methyl-5-oxo-
1H—1,2,4-triazol-1-
yl]phenyl]methanesulfonamide

N-[[(4,6-dimethoxy—2-
pyrimidinyl)amino]carbonyl]~2-
(ethylsulfonyl)imidazo[ 1 ,2-a]pyridine-3-
sulfonamide

 

130

mgrmmrlggnjiﬂrm1111;an