Published December 13, 2018
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Hail Damage Impacts on Corn Productivity:
A Review
Martín Battaglia,* Chad Lee, Wade Thomason, John Fike, and Amir Sadeghpour
ABSTRACT
Hail influence on corn (Zea mays L.) yield depends
on defoliation timing and severity. Complete
defoliation during early vegetative stages can
have minimum yield effects if plants’ growing
point is not affected but can generate some
delays in the planting to flowering period. Low-
severity defoliations after V10 can reduce yield
up to 30%. Higher severities gradually increase
yield penalties to a peak around flowering and
decrease progressively during the grain-filling
period. Charts to estimate the percentage of
corn yield loss due to defoliation developed in
the late 1960s are still accurate in most situa-
tions but fail to describe particular situations.
Defoliation around VT commonly affects time
to silking, anthesis–silking interval, and plant
growth rate, but not time to anthesis, and is
commonly explained by lower kernel number
(KN). Defoliation at R2 commonly affects kernel
weight (KW), without changing KN. However,
several studies showed a reduction in both KW
and KN with R2 defoliations. Under low plant
disease pressure, fungicides applied around
VT do not help reduce any yield defoliation
impact. Specific genotypes, row spacing, and
hybrid maturity can influence crop yield defolia-
tion responses. More studies are warranted to
confirm the potential for narrow rows to reduce
yield loss after defoliation.
crop science, vol. 59, january–february 2019  www.crops.org 1
REVIEW & INTERPRETATION
M. Battaglia, W. Thomason, and J. Fike, Dep. of Crop & Soil
Environmental Sciences, College of Agriculture and Life Sciences,
Virginia Tech, Blacksburg, VA 24061; C. Lee, Dep. of Plant and
Soil Sciences, College of Agriculture, Food and Environment, Univ.
of Kentucky, Lexington, KY 40546-0312; A. Sadeghpour, Dep. of
Plant, Soil and Agricultural Systems, College of Agricultural Sciences,
Southern Illinois Univ., Carbondale, IL 62901. Received 3 May 2018.
Accepted 22 Oct. 2018. *Corresponding author (martinb1@vt.edu).
Assigned to Associate Editor Lucas Borrás.
Abbreviations: ASI, anthesis–silking interval; DON, deoxynivalenol;
IPAR, intercepted photosynthetically active radiation; KN, kernel
number; KW, kernel weight; LAI, leaf area index; PGR, plant growth
rate; QoI, quinone outside inhibitor; RM, relative maturity; RI,
radiation interception; ZEA, zearalenone.
H
ail stones, irregular ice clumps created in convective clouds
(Changnon and Kunkel, 2003), can cause significant damage
to crops and property. As a weather phenomenon, hail storms are
highly variable both temporally and spatially, but these events
have produced large economic losses for US agriculture, particu-
larly in corn (Zea mays L.) crops. The nation’s areas of greatest
hail frequency are along and just east of the central Rocky Moun-
tains, which typically experiences 6 to 12 hail events per year
(Changnon et al., 2009). The valleys of the Rocky Mountains
have the nation’s greatest hail intensity with the largest average
stone sizes, whereas the lowest intensities are found in the eastern
and southwestern United States, where hail storms occur once
every 2 or 3 yr (Changnon et al., 2009). In the continental United
States, approximately half of all hailstorms occur between March
and May (Klein and Shapiro, 2011), and about a third of all hail-
storms occur between June and September, a time period when
replanting options are not economically sound for farmers (Vorst,
1993). Overall, the average annual frequency of crop-damaging
hail across the United States is 158 d (Changnon et al., 2009).
Published in Crop Sci. 59:1–14 (2019).
doi: 10.2135/cropsci2018.04.0285
© Crop Science Society of America | 5585 Guilford Rd., Madison, WI 53711 USA
All rights reserved.

Long-term trends (1948–1997) indicate increased
frequency of hail events in the High Plains and Southeast
and decreases in the Midwest and West (Changnon, 2000).
However, nationwide, hail events and crop and property
losses from hail all show downward trends for the period
of 1950 to 2009 (Changnon et al., 2009). Total hail losses
for all crops were estimated in the late 1990s by Changnon
(1997) at about US$1.3 billion annually, representing
between 1 and 2% of the annual crop value. Hail losses
vary considerably regionally, representing, for example, 1
to 2% of the crop value in the Midwest, 5 to 6% of the
crops produced in the High Plains, and much less elsewhere
in the nation (Changnon, 1997). Corn grain yield losses
due to hail damage cost US producers about $580 million
annually (Changnon et al., 2009). Unusual years with high
hail incidence like 2014, when crop insurance across the
United States registered the largest economic losses ever,
may increase this value several times; however, for example,
a single hail event on 3 June 2014 caused injured losses of
approximately $150 million in the state of Nebraska (http://
www.air-worldwide.com/Models/Agriculture/).
Understanding the impacts of hail damage in corn plant
stands and the best strategies to manage them is important
to minimize the economic losses produced by hail damage.
The objective of this review is to summarize the current
knowledge in relation to the documented effects of hail on:
(i) corn grain yields before and after the V10 growth stage,
(ii) corn grain yield components, (iii) dynamics of anthesis
and silking processes, and (iv) plant disease pressure across
the United States over the last 200 yr.
TYPES OF HAIL DAMAGE
IN CORN PLANTS
Corn plants damage by hail mainly result in loss of photo-
synthetic area. Hail most often produces grain yield losses
by shredding leaf blades, resulting in a loss of leaf area
(Dungan, 1934; Jenkins, 1941; Hanway, 1969; Vorst,
1993; Klein and Shapiro, 2011). In severe cases, the entire
leaf blade is stripped from the leaf midrib. Other damage
to corn plants occurs when hail strikes small plants,
breaking them at the soil surface and reducing plant
stands (Dungan, 1934; Vorst, 1993; Nielsen, 2012). Hail-
stones also bruise the stalks of larger plants, resulting in
interference with the movement of assimilates (Dungan,
1934) and increasing chances of plant breakage by winds
(Eldredge, 1935). If hail occurs at pollination, it may seri-
ously interfere with normal fertilization of the kernels
through the destruction of the functioning organs of
the tassels and silks (Dungan, 1934). Storms later in the
season may result in direct damage to the developing ears,
causing complete ear loss or reducing the quality of the
surviving kernels bruised by the hail stones. According
to Dungan (1934), the diversity of types and degrees of
injury that can be inflicted by hailstorms makes it almost
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impossible to duplicate in experimental research. Even
more, the severity of a hailstorm typically varies spatially,
even in a small field. Nonetheless, manual defoliation—
removing a given percentage of the leaf area at different
moments in the crop cycle—can be an effective research
tool for understanding the effects of hail damage on grain
yield loss (Dungan, 1934).
RESEARCH ON METHODS AND TYPES
OF LEAF REMOVAL
The potential photosynthetic contribution of leaves
located at different heights within the crop canopy has
been studied in defoliation experiments. Several studies
with older hybrids indicated that leaves above and below
the ear were equally critical to grain yield (Hanway, 1969;
Egharevba et al., 1976). However, the relative contri-
bution of upper leaves to overall plant productivity in
modern corn hybrids may be more important than for
older hybrids (Adee et al., 2005; Subedi and Ma, 2005).
Hanway (1969) compared six leaf removal methods
(50% removal) at VT and R2 stages using a midseason
hybrid. Removing (i) alternate pairs of leaves (every other
two leaves), (ii) all leaves from one side of each plant,
(iii) half of each leaf (lengthwise), (iv) all leaves above the
uppermost ear, (v) all leaves below the uppermost ear, or
(vi) the terminal half of each leaf reduced grain yields 17
to 23%, but differences among treatments were not signifi-
cant. Similarly, Egharevba et al. (1976) did not find grain
yield difference when comparing complete defoliation
from below and above the uppermost ear within 30 d of the
R1 stage and concluded that leaves both above and below
the ear were critical for final grain yields. Egharevba et al.
(1976) hypothesized that this might be a mixed response
resulting from a decreased ability to fix CO2 over time
for the lower leaves, and the fact that mid- and late-season
hybrids have ?60% of total leaf surface under the main
ear. Thus, larger leaf area below the ear may have compen-
sated for the reduced photosynthetic efficiency associated
with leaf age. In contrast, only 12% reductions in final grain
yield were measured when all the leaves below the ear were
removed, whereas 88% yield reduction was measured with
the removal of all leaves above the ear at R1 in a more
recent study conducted in Illinois (Adee et al., 2005). Simi-
larly, removal of all leaves above the ear at the VT growth
stage reduced plant dry matter 2.3 and 12.3 times more
than removal of all leaves below the ear in a conventional
grain hybrid and a leafy silage corn hybrid in Canada,
respectively (Subedi and Ma, 2005).
HAIL DAMAGE IMPACTS ON CORN
GRAIN YIELD
Much research has been undertaken to understand the rela-
tionship between defoliation timing, defoliation severity,
and final grain yield in corn. The first field experiments

conducted in the United States focused on the study of
the practice named “topping.” This practice, common in
the United States in the 1800s and early 1900s, consisted
of cutting the corn plant right above the insertion of the
main ear around the dent stage (i.e., R5; Abendroth et
al., 2011) to use for livestock feeding (Dungan, 1934).
Occasionally, this was followed by a practice known as
“stripping,” which implied the removal of all leaf blades
below the main ear (Dungan, 1934). With a few excep-
tions (e.g., Buel, 1842), these practices were resisted by
scientists during the first half of the 19th century. Ruffin
(1835) defined them as “among the most irrational prac-
tices in agricultural pursuits” and added that “at not
distant day farmers will condemn the present practice.”
Tucker (1837) claimed that “among enlightened farmers,
the practice of topping has fallen into general disrepute,
and as it is most clearly unprofitable practice, it should
be entirely abandoned.” Tracy and Lloyd (1895), summa-
rizing the work on the impacts of corn topping from eight
research stations across the United States in the late 1800s,
reported grain yields losses of 16 to 18% after topping
corn stands in comparison with control plots. Moreover,
these reductions in yields were not compensated by the
extra value of the forage obtained for livestock feeding
(Connell, 1890; Hunt, 1891).
In general, grain yield losses from hail damage to
young plants, up to 3 wk postemergence, is often minimal
( Jenkins, 1941) because the growing point is still below
the surface, thus enabling high tolerance to hail damage
(Vorst, 1993; Lee, 2007). From this time and approxi-
mately until to V9 to V10 growth stages, defoliation
treatments imposed to simulate hail damage reduced corn
grain yield in several studies (Eldredge, 1935; Lindstrom,
1935; Dungan and Gausman, 1951; Cloninger et al., 1974;
Johnson, 1978) but did not change it (Crookston and
Hicks, 1978; Andrade et al., 2002; Battaglia et al., 2018)
or even increase it in others (Crookston and Hicks, 1978).
After V10, plant nutrient and dry weight accumulation,
and therefore the overall water and nutrient requirements,
greatly increase in corn (Ritchie et al., 1993; Abendroth
et al., 2011), and the plant becomes more vulnerable to
hail damage (Vorst, 1993). As a result of this, damage
increases gradually to a peak in the period just preceding
or a few days after tasseling, when complete defoliation
can cause grain yield losses as high as 100% (Hume and
Franzke, 1929; Eldredge, 1935; Hanway, 1969; Egharevba
et al., 1976; Vasilas and Seif, 1985a; Andrade et al., 1999;
Adee et al., 2005). Grain yield losses to defoliation after
tasseling decrease progressively approaching maturity
(Vorst, 1993), although they can still reach values of up
to 40% with complete defoliation at R3 (Eldredge, 1935)
or R4 (Andrade, personal communication, 2012), even-
tually reaching 0 (Dungan, 1934; Jenkins, 1941; Adee et
al., 2005; Battaglia et al., 2018) to 5% (Eldredge, 1935) at
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maturity. At the same time, most studies have consistently
shown that greater grain yield losses occur with 100%
defoliation than with 50% defoliation (Dungan, 1934;
Hanway, 1969; Hicks et al., 1977).
Response to Corn Defoliation before the
V10 Growth Stage
Dungan (1934) suggested that treatments earlier than the
VT growth stage are not so detrimental to grain yield for
two reasons: (i) some new leaves may still unroll with the
elongation of the tassel-bearing stem, and (ii) after defolia-
tion, the photosynthetic efficiency of the remaining leaves
is increased by the removal of blades, thus preventing
the grain yields from falling off in direct proportion to
the amount of leaf area destroyed. However, complete or
near-complete defoliation on plants before V6 resulted
in differing results in other studies. Lindstrom (1935)
studied the grain yield response after removing 75 to 85%
of the upper aerial biomass (wet weight basis) in six F1
inbred progenies on four different dates, starting at 9 d
and finishing at 21 d after emergence. Overall, grain yield
decreased for all the defoliation dates, with reductions
ranging from 15 to 33% on plants defoliated at 9 and 21
d after emergence, respectively (Lindstrom, 1935). Clon-
inger et al. (1974), working in Missouri with 28 crosses
among eight inbred lines, documented that cutting V3,
V4 to V5, and V6 corn plants at the ground level decreased
grain yields by 11, 38, and 46%, respectively. Similarly,
complete removal of aboveground biomass between V3
and V6 growth stages decreased yields 5 to 44% in other
studies (Eldredge, 1935; Dungan and Gausman, 1951).
Dungan and Gausman (1951) observed that cutting V7
to V8 plants at the soil level has the potential to produce
almost complete grain yield loss (i.e., 97%) in some cases,
whereas cutting the same plants at 5 cm above ground level
can reduce those losses by up to 54%. The authors attrib-
uted the latter response to the fact that the growing point
in plants at the V7 to V8 stage were within 5 cm of ground
level, precluding its removal by the imposed treatments.
Corn relative maturity (RM) group may play an
important role in responses to defoliation in some cases
(Crookston and Hicks, 1978), but not in others ( Johnson,
1978; Andrade et al., 2002). Crookston and Hicks (1978)
reported that complete defoliation at V3 to V4 in early-
maturing hybrids may boost final grain yields. Working in
Minnesota with 12 different early-maturing hybrids (70 to
90 RM) in 75-cm row widths, they observed that complete
defoliation at V3 to V4 caused an average 13% grain yield
increase in one location, and a range of responses ranging
from no change up to 22% yield increase across three other
locations. In another experiment, a side-to-side compar-
ison between an early (90 RM) and full-maturity hybrid
(115 RM) resulted in a 56% increase and 7% grain yield
decrease for complete defoliation at V3 to V4 compared

with the undefoliated control, respectively (Crookston
and Hicks, 1978). Authors hypothesized that the V3 to
V4 defoliation occurred at or near floral initiation for the
early, but not for the full-maturity hybrid. A sudden loss
of vegetative tissue in a plant that is developing vegetative
and reproductive structures would likely induce a drastic
change in source–sink relations, in turn stimulating
embryonic ear growth at the time of reproductive initia-
tion. In contrast, Johnson (1978) observed that, regardless
of maturity, complete defoliation of several early-, mid-,
and full-season hybrids at V2, V3, and V4 reduced final
grain yield by an average of 12% and leaf area index
(LAI) by 22% at two Illinois locations over a 2-yr period.
Johnson (1978) concluded that there was no evidence that
grain yields and loss in LAI in early-season hybrids were
less sensitive to defoliation than yields and LAI of full-
season hybrids. In addition, defoliation from V2 to V4 in
this study did not have a beneficial effect on the grain yield
of corn. Similar trends were documented by Andrade et
al. (2002), who did not find differences in yield between
a mid- and a full-season corn completely defoliated at V3
and V5 in either 52- and 70-cm rows working in Argen-
tina. Maximum grain yield was achieved for both hybrids
and row widths with defoliations at V3. Yields did not
differ from control treatments, as corn plants reached at
least 80% maximum solar radiation interception (RI) at
silking (versus at least 85% RI in the control). Grain yields
for both hybrids were similarly reduced with defoliation at
V5, but reductions were greater with 70-cm (−20%) than
with 52-cm (−9%) row spacing; plants intercepted only 63
and 73% RI at silking in each case, respectively.
Response to Corn Defoliation after the
V10 Growth Stage
Dungan (1934) removed 8, 17, 25, 33, 50, 67, 83, and
100% of leaves at the tasseling (VT), fresh silking (R1),
early blister (R2), milk (R3), and early dent stages (R5).
Plant development stage had little influence on grain yield
reduction with up to 25% leaf removal. Grain yield loss
reached 100% for 100% blade removal at R1 and dropped
gradually with subsequent growth stage, reaching 0% loss
at R5. These findings supported results from previous
work from the late 1800s to early 1930s (Connell,
1890; Tracy and Lloyd, 1895; Hume and Franzke, 1929;
Culpepper and Magoon, 1930; Dungan, 1930).
The early generation of leaves and their posterior
maintenance close to or during the critical period in corn
(i.e., ±15–20 d bracketing VT to R1 stages) are impor-
tant variables affecting the light interception efficiency
(Williams et al., 1965; Maddonni and Otegui, 1996;
Cárcova et al., 2003a, 2003b) and have profound impacts
on the final grain yield achieved (Andrade et al., 1993).
Theoretical increases in LAI of up to 6 to 6.5 increase
intercepted photosynthetically active radiation (IPAR) to
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values between 90 and 95% (Williams et al., 1965; Gallo
and Daughtry, 1986). However, these LAI values are rarely
measured in the field. Bradley and Ames (2010) measured
greater LAI at V12 in undefoliated corn plants than in
corn plants defoliated between 25 and 30% (LAI values of
3.2 vs. 2.3, respectively). In this experiment, defoliation
rates of 30% or less at the V12 stage were enough to reduce
yields by 18 to 30%. In a 2-yr experiment conducted in
Balcarce, Argentina, Andrade (personal communication,
2012) studied the effect of four levels of defoliation (i.e.,
control [0%], mild [33%], moderate [50%], and severe
[100%]) imposed at V17, R1, and R4 on corn grain yield.
Control plants reached 95 and 80% IPAR for irrigated and
rainfed conditions, respectively. Average IPAR for mild
(33%) and moderate (50%) defoliation was ?80%, being
always slightly lower for 50% than for 33% defoliation.
With severe defoliation, crops did not exceed 40% IPAR.
Grain yield was reduced 94% with 100% defoliation at
V17 and R1 and 42% with 100% defoliation at R4. Mild
and moderate defoliations decreased grain yields by 13%
with no differences due to timing of defoliation.
Contrasting grain yield responses to defoliation for
different corn RM were more noticeable with treat-
ments imposed immediately before or after VT in the
studies conducted by Hicks et al. (1977). Defoliation
of an early (90 RM) and full-season (115 RM) hybrids
reduced average grain yield by 30, 100, and 45% for
complete defoliations (i.e., 100%) at V11, VT, and R3,
respectively. Complete defoliation at R5 did not affect
yield for the 90-RM hybrid but decreased yield by 12%
in the 115-RM hybrid. Partial defoliation (i.e., 50%) at
V11 produced similar grain yield reductions of 5% for
both hybrids. However, partial defoliation at the VT stage
reduced grain yields less in the 90-RM (11%) than in the
115-RM hybrid (30%). Partial defoliation during R3 and
R5 increased yield by 3.5 and 13% in the 90-RM hybrid,
respectively, but reduced yield of the 115-RM hybrid by
9 and 5%, respectively. These responses may be explained
by the improved water use efficiency of partially defoli-
ated short-season plants exposed to low soil water contents
at reproductive stages in at least 1 of the 3 yr under study,
and not because of differences in post-defoliation IPAR
or LAI between the two hybrids (Hicks et al., 1977).
Conversely, Hanway (1969) found that defoliation caused
greater grain yield losses for an early- than for a mid- and
late-season hybrid in Iowa. Defoliation at the V10, VT,
and R2 stages reduced average grain yield by 10 to 19,
17 to 28, and 17 to 22%, respectively, for 50% defoliation
and by 17 to 42, 97 to 99, and 66 to 74%, respectively,
for 100% defoliation. In this study, three different plant
populations had no significant effect on grain yield loss as
a result of defoliation treatments. In the other extreme of
responses, Egharevba et al. (1976) reported no significant
interaction between hybrids and grain yield for defoliation
 14350653, 2019, 1, Downloaded from https://acsess.onlinelibrary.wiley.com/doi/10.2135/cropsci2018.04.0285 by Purdue University (West Lafayette), Wiley Online Library on [29/11/2022]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
initiated at R1 and continued every 10 d for the next

100%
—————————————————————————————————————————————————— % ——————————————————————————————————————————————————
40 d in an early-, mid-, and late-season hybrid in a

59
41
9
11
13
16
22
28
34

84
96
100
97
73
44
51
61
72

23
0
2-yr study in Missouri.

95%
Almost all experiments available in the litera-

54
38
87
9
10

91
88
66
12
15
20
26
31
40
46
55
65
76

21
0
ture were designed to show the effect of defoliation

90%
treatments on the corn grain yield at a specific stage.

35
49
83
80
60
8
9
11
14
18
23
28
36
42
49
59
69
79

20
0
Interestingly, Adee et al. (2005) used the term “corn
defoliation progress curves” in a defoliation experi-

85%

32
45
7
8
10
13
16
20
25

75
72
55
32
38
44
53
62
71

19
0
ment in northern Illinois to describe corn response to
progressive defoliation initiated at V14 through R5.

80%
Grain yield (in units of Mg ha−1) declined linearly

29
41
6
7
9
11
14
18
22

68
65
50
28
34
40
48
56
64

17
0
with defoliation initiated at V14 through R3, but the

75%
percentage of grain yield loss associated with any level

26
37
5
6
7
9

62
58
45
12
16
19
25
30
36
43
50
57

15
0
of defoliation did not differ between V14 and R3. Defo-
liation after the R3 stage resulted in a lesser grain yield

70%

32
23
51
5
6
7
9
11
15
17

55
51
39
22
26
31
37
44

14
0
loss, with no change in yields from R5 defoliation.

65%
Percentage leaf area destroyed

28
20
4
5
6
8
10
13
15

48
45
34
20
23
27
32
38
43

12
0
GRAIN YIELD RESPONSE TO
SIMULATED HAIL DAMAGE IN FIELD

60%

24
17
4
5
6
8
9
11
13
17

28
33
38
42
39
30

10
20
23

0
STUDIES VS. CHARTS USED BY
INSURANCE COMPANIES 55%

21
14
3
4
5
7
8
10
11
15

32
36
34
26
17
20
24
28

8
0
Hail insurance companies have used the National
Table 1. Estimated yield loss percentage as a result of corn defoliation at different growth stages.†

Crop Insurance Service’s charts (Table 1) to estimate

50%

the percentage of corn yield loss due to defoliation

18
12
2
3
4
6
7
9
10
13

27
31
29
22
15
18
21
24

7
0
at various stages since the late 1960s in the United

† Source: adapted from the National Crop Insurance Association’s “Corn Loss Instructions” (revised 1984) in Vorst (1993).
45%

States (Lee, personal communication, 2013). At the

15
10
1
2
3
5
6
7
8
10

22
26
24
19
12
14
17
19

6
0
field level, crop adjustors use the “hail adjustor’s hori-
zontal leaf method” to quickly determine the corn
40%

8
1
1
2
4
5
5
6
8

18
21
20
16
12
9
11
13
15

4
0
growth stage when hail damage has occurred. Vege-
tative stages in this scale are determined by counting
35%

the number of leaves tips that are bending over.

6
0
1
2
3
3
4
4
6

14
17
16
13
9
7
8
9
11

3
0
However, this method does not account for the first
30%

oval-shaped leaf, and it usually needs some approxi-

4
0
0
1
2
2
3
3
4

11
13

10
7
5
6
7
9

12

2
0
mation (i.e., splitting a lower corn stalk, and locating
25%

the first node, usually connected to the fifth leaf ) to

2
0
0
1
1
1
2
2
3

8
9
9
7
5
3
4
5
7

1
0
determine growth stage beyond the eight-leaf stage
when leaves usually slough off from the corn stalk,
20%

2
0
0
0
0
1
1
1
2

6
7
7
5
3
2
3
4
5

0
or in situations where upper leaves are missed after a 0
severe hail storm (Lee, 2007; Thomison, 2016). Crop
15%

extension agronomists and professional agronomists

1
0
0
0
0
0
0
1
1

4
5
5
3
2
1
2
3
3

0
0

have widely adopted the “leaf collar method,” in

10%

which leaves with visible collar are counted as fully

0
1
0
0
0
0
0
0
0
0

3
3
3
2
1
1
1
2
2

developed leaves. Wide adoption of this method, at

both the research and extension levels, and its greater
Approx. equivalence
with leaf collar

degree of accuracy in determining corn vegetative

‡ Adapted from Abendroth et al. (2011).
method‡

stages even after severe hail storms may be important

V17–V19
V10

V16
V13

V15
V12

V14
V11

R5
R2
R3
R4

R6
VT
R1
V7

V9
V8
V5
V6

features to determine its adoption by crop insurance

adjusters in the near future.
Currently, losses resulting from corn defoliation
due to hail damage prior to V5 (collar method) are
19- to 21-leaf

calculated based on the need or lack of need to replant

Soft dough
Stage of

corn stands, and not as a percentage of the final grain

growth

Mature
10-leaf

16-leaf

18-leaf
13-leaf

15-leaf
14-leaf
12-leaf
11-leaf

17-leaf

Tassel

Blister
Silked
9-leaf
8-leaf
7-leaf

yield loss (Table 1). However, research conducted in

Dent
Milk

crop science, vol. 59, january–february 2019  www.crops.org 5


the 1970s in Illinois, Minnesota, and Missouri showed
that defoliations between V2 and V4 had the potential to
produce both negative and positive grain yield responses
ranging between −8 and +13% in most comparisons.
In some particular situations, yields were reduced up to
22% and grain yield increased up to 48% (Cloninger et
al., 1974; Hicks et al., 1977; Crookston and Hicks, 1978;
Johnson, 1978) (Table 2). Clearly, these potential boosts
in final grain yield after early defoliations in corn are
not currently accounted for at the time farmers or crop
adjusters evaluate replanting options.
An ample majority of the grain yield data from defolia-
tions performed between the V5 and V9 stages in previous
studies (Eldredge, 1935; Dungan and Gausman, 1951;
Cloninger et al., 1974) are not comparable with the esti-
mated grain yield loss for a similar growth stage in Table
2. In the first case, most defoliations were performed at
heights either at or around 5 to 6 cm above ground level,
resulting in grain yield losses of up to 97% with ground-
level defoliations at V8. These situations, where most of
the plant is damaged from planting up to V8, are more
properly covered by the charts used to evaluate replanting.
When situations were comparable (i.e., only the laminar
part of the leaf was defoliated), complete defoliations at V5
resulted in grain yield losses of 19% in Ohio (Mangen et
al., 2005), compared with chart values of 9% for a similar
comparison (Table 1). Battaglia et al. (2018) found no
evidence of grain yield loss after complete defoliation at
the V7 stage in most comparisons in Kentucky, compared
with chart yield losses between 11 and 13% for defoliations
>90% at the V7 stage (Table 1).
Chart grain yield losses for partial and complete defo-
liations at the V10 and V11 stages (9–10 and 28–34%,
Table 1) are similar to those reported by Hanway (1969)
in Iowa and Hicks et al. (1977) in Minnesota for partial
(5–15%) and complete defoliations (30%) at the V10 to V11
stage, respectively (Table 2). Partial defoliations at the V12
(Vasilas and Seif, 1985b) and V13 (Mangen et al., 2005)
stages resulted in grain yield losses between 8 and 15%,
which is comparable with chart losses between 13 and
15% for identical comparisons (Table 1). However, chart
losses used by insurance companies may not be adequate
to assess the impact of complete defoliation occurring
immediately before the occurrence of the VT to R1 stage
in inbreds lines or specialty crops. In the V12 stage, 100%
defoliations performed by Vasilas and Seif (1985b) led to
yield losses across six contrasting inbreds ranging from 25
to 74% (Table 2), compared with a hail insurance grain
yield loss of 44% (Table 1). In the case of Mangen et al.
(2005), yields for high- (91% male-sterile hybrid and 9%
pollinator seed) and normal-oil hybrids (100% male fertile
plants) declined between 28 and 30% after 100% defolia-
tions at V13, compared with chart values of 51% (Table 1).
Data on the impact of corn defoliation after the VT to R1
6 www.crops.org crop science, vol. 59, january–february 2019
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critical period in corn are not available for either inbreds
or specialty crops.
The differences between values used by crop adjusters
and those reported in field experiments tend to disappear
with defoliations occurring immediately before or after
the VT to R1 stage in corn hybrids for grain production.
Hanway (1969) observed grain yield losses of 25 and 98%
after partial and complete defoliations at VT, compared
with values of 31 and 100% for identical comparisons in
Table 1. Similarly, other authors have reported grain yield
losses close to or at 100% for complete defoliations at VT
or R1 stages (Eldredge, 1935; Hanway, 1969; Egharevba
et al., 1976; Adee et al., 2005; Battaglia et al., 2018). It is
evident that genotypic differences in their yield response
to defoliations exist (Tamagno et al., 2016).
Battaglia et al. (2018) reported grain yield losses
between 4 and 27% for partial and >85% for complete
defoliations at the V14 stage, compared with values of 18%
for 50% defoliation at V14 and 61% grain yield loss for
100% defoliation at V14 in Table 1. Over the 2-yr study
in Kentucky, V14 occurred 2 to 4 d before the VT stage,
which explains the similarity between the values reported
in this experiment and the grain yield losses around the
VT stage in Table 1. However, analysis of the data shows
a significant triple interaction among hybrids (113 and
120 RM), rows (38 and 76 cm), and defoliations in 1 yr,
with 50% defoliations at V14 reducing the grain yield of
the 120-RM hybrid in narrow rows (−27%) more than in
wide rows (10%), and vice versa (113-RM hybrid grain
yield was less reduced in narrow [−4%] than in wide rows
[−27%]). Similar interactions were observed for 50% defo-
liations at R2 in this study. Complete defoliations at the
VT or R2 stage, regardless of the levels of other factors,
reduced grain yields by >85% in most comparisons.
CORN GRAIN YIELD COMPONENTS
AFFECTED BY DEFOLIATION
Kernel number (KN) per surface area and kernel weight
(KW) are the main yield components reported for most
crops (Egli, 1998). In the case of corn, the number of
kernels set per plant is highly dependent on the environ-
ment around silking (Kiniry and Ritchie, 1985; Tollenaar
et al., 1992; Andrade et al., 1993; Kiniry and Knievel,
1995). Consequently, plant physiological status around this
stage is critical for final KN in corn (Andrade et al., 1999),
since corn is highly inefficient in the use of assimilates
stored before silking (Kiniry et al., 1992; Borrás et al.,
2004). Kernel weight is also an important component of
final grain yield and is highly dependent on the postsilking
environment (Cirilo and Andrade, 1996; Maddonni et al.,
1998; Borrás et al., 2004).
Grain yield losses associated with defoliation around VT
or R1 (tasseling or silking) were mainly explained by fewer
KN in numerous studies (Culpepper and Magoon, 1930;
 Table 2. Summary for relevant defoliation studies.
Reference Study Hybrids or inbred lines† Maturity‡ Defoliation timing§ Defoliation severity¶ Final yield# Yield component affected††
Dungan (1934) 1925–1929 1925–1928: uniform strain of – VT, R1, R2, R3, and R5 8.3, 16., 25, 33.3, 50, 66.7, ?100% loss for 100% defoliation at VT and R1; No reference about KN; KW most
(Illinois) open-pollinated Reid Yellow Dent 83.3, and 100% then decreasing until reaching 0% at maturity; reduced by R2 defoliation
little effect of defoliation up to and below 25% at
1929: F 1365 any stage other than VT
Lindstrom (1935) 1933 6 lots of F1 inbred progenies – From +9 to +21 d after 75 to 85% top plant aerial 15 to 33% yield loss from early to late –
(Iowa) planting portion cutting, respectively

Dungan and 1945–1948 1945–1947: double cross-hybrids – V6 and V7–V8 Cutting at soil level and 5 V6: 44 and 26%; V7–V8: 97 and 54% –
Gausman (1951) U.S. 13, Illinois 201, and Illinois cm above soil level at each yield losses for ground and +5-cm level,
(Illinois) 972 stage respectively

1948: eight adapted single crosses – V5–V6 and V8–V9 Cutting at soil level and 6 From 18 (V5–V6, soil level) to 48% (V8–V9, 6 –
and eight standard inbreeds cm above soil level cm above soil) yield reduction

Hanway (1969) 1965 3 hybrids: early (B8xW153R), – V10, VT, and R2 50 and 100% 15, 25, and 20% for 50% defoliation and 30, KN and KW for all treatments except
(Iowa) midseason (Wf9xB37), and late 98, and 69% for 100% defoliation at V10, VT, 100% at VT (complete barrenness)
(B14xC131A) and R2, respectively

Cloninger et al. 1969–1970 28 single crosses among 8 74 to 88 d to VT V3 to V4–V5 to V6 Ground-level cutting Average 11, 38, and 46% yield reduction for –
(1974) (Missouri) inbreeds lines (Va35, H49, B57, V3, V4–V5, and V6, respectively; yield increase

crop science, vol. 59, january–february 2019 

B37, B14A, Mo17, Mo5, N38A) in five singles crosses clipped at V3 and one
cross clipped at V4–V5

Egharevba et al. 1971–1972 P3773, P3306, and P3149 Early-, mid-, and Starting 10 d after R1 and All leaves below ear, all Complete defoliations more detrimental KN most reduced with 100%
(1976) (Missouri) late-season, continued at 10-d intervals leaves above ear, and all (6–82% losses) than partial defoliation defoliation from +0 to +10 d after 50%
respectively for the next 40 d leaves (2–33% losses) R1; KW most important from +20
d and more from 50% at R1 and/or
partial defoliation

Hicks et al. (1977) 1973–1975 Short-season (Trojan TXS 85) and 90 and 115 RM V3–V4, V11, VT, R3, and 100% at V3–V4; 50 and 100% yield loss for 100% defoliation at VT; KN not recorded; KW not affected
(Minnesota) full-season DeKalb XL45a R5 100% for the rest slightly greater yield losses at R2 (range = +4 before VT; after VT, KW was greatly
to −47%) than V11 defoliation timing (range = reduced only at 100% defoliation
−3.5 to −31%); 100% rates always produced rates
greater losses than 50% rates

Crookston and 1973–1975 Short-season (Trojan TXS 85) 90 and 115 RM V3–V4 100% 8% yield loss for full-season hybrid; 48% KW not affected; authors hypothesized
Hicks (1978) and full-season DeKalb XL45a average increase for short season KN was completely affected
(Minnesota)
1975 12 short-season hybrids 70–95 RM V3–V4 100% Average yield increase of 13% (range = +37 –
to −14%)

1976 12 short-season hybrids (three 70–95 RM V3–V4 100% Average yield response ranged from 0 to –
locations) −22% from southern to northern locations
(Minnesota)
Johnson (1978) 1976–1977 9 hybrids (Funks 5048, Trojan 8 early-season V3–V4 100% 11.2% yield decrease (averaged across both Smaller ears
(Illinois) TXS85, Sokota SK36, Pioneer hybrids; DeKalb years and 9 hybrids)
3976, Jacques JX62, DeKalb XL43a full-season
XL12, DeKalb XL310, Minhybrid
7301, DeKalb XL43a)

1976 4 early-season and one full- – V3–V4 100% 13% yield decrease (averaged across 5 Smaller ears
season hybrids hybrids)

1977 2 early-, 2 mid-, and 2 full- – V2, V3, and V4 100% 10% yield decrease (averaged across 6 Smaller ears
season hybrids hybrids)

www.crops.org 7
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 8
Table 2. Continued.
Reference Study Hybrids or inbred lines† Maturity‡ Defoliation timing§ Defoliation severity¶ Final yield# Yield component affected††
Tollenaar and 1970–1971 1970: United 106, 1971: United – 1970: 0, 2, 4, and 6 wk 50 and 100%, only 100% Maximum yield for control 6 wk after R1: 15% KN: 100, 61, and 16% loss with
Daynard (1978) 106 and Warwick SL 209 after R1, 1971: 1 wk after for the extra treatment yield loss for 50 and 100% defoliation; 0, 2, 100% defoliation after 0, 2, and 4 wk
(Ontario) R1 added included in 1971 and 4 wk after R1: 21, 29, and 38% loss for from R1; 50% defoliation produced
50% severity; 46, 73, and 100% for 100% losses of up to 26% only within first
severity; 1 wk after R1 (1971): 89% loss 2 wk from R1; KW: not affected
with defoliation within 4 wk from R1;
reduction of up to 38% in KW with
100% defoliation after 4 wk from R1

Vasilas and Seif 1982–1983 6 inbreds (FR16, MS17, – V12.5 50 and 100% 100% defoliation: greatest yield loss (−74%) for Yield reductions explained more
(1985b) (Illinois) FRM017rbm, FR25, FR632, and FR16, FRM017rbm, and FR632; intermediate by reductions in KN (6 to 58% with
FR27rbm) loss (−54 to −57%) for MS17 and FR25, and complete defoliation; <10% with partial
lowest (−25%) for FR27rbm; 50% defoliation: defoliation); only 100% defoliation
yield loss ranging from 8 to 13%, with +9% reduced KW, but <10% in most
yield increase in inbreds FR16 and FR27rbm comparisons

Yao et al. (1991) 1988–1989 Ferke 7928 – One defoliation in 1988; 100% All defoliation reduced grain yield compared Control (T1) had ?6% greater KN per
(Ivory Coast) control (T1), two (T2), four with the control (T1), with yield losses cob than T2 and T3; defoliation delayed
top (T3), all leaves above ranging from 46 (T4) to 100% (T2) silking but not tasseling; all defoliation
(T4), or below ear (T5), or reduced KN (range = −32 to −100%)
three top leaves above ear and KW (range = −55 to −100%)
undefoliated (T6)

Adee et al. (2005) 1997–1999 P3394, P3489, and P33Y18 2660, 2630, and V14, R1, R2, R3, and R5 20% of the leaf area at Yield declined linearly with defoliation initiated –
(Illinois) 2710 GDD one or more corn growth at V14 through R3; losses associated with
stages any level of defoliation did not differ between
V14 and R3; after R3, yield losses diminished;
defoliation at R5 did not affect final yield

Mangen et al. 1999–2000 Two high-oil corn (HOC) TopCross NOC: 108 RM V5 and V13 50% (only V13), 100% (V5 100% defoliation at V5 and 50% defoliation KN was reduced between 7 and 11%

www.crops.org
(2005) (Ohio) blend (P34K79 and P34B25) and
two normal-oil corn (NOC) hybrids
(P34K77 and P34B23)
and 1480 GDD and V13) at V13 reduced yield only in 2000 (−19 and by 100% defoliation at V5 and 100%
(P34K77); 12%); 100% defoliation at V13 reduced defoliation at V13 only in 2000; 50%
109 RM and 1500 yields by 28–30% (no differences in yield defoliation at V13 did not affect KN;
GDD (P34B23) between HOC and NOC hybrids; no hybrid KW was reduced 7 to 11% by 100%
´ defoliation interaction for all comparisons) defoliation at V13

Echarte et al. 2000–2001 DKF880, M400, DK4F36, DK664, 120, 128, 127, 116, 27 d (?330 GDD) after Aiming at reducing canopy – Defoliation 27 d after R1 reduced KW
(2006) (Argentina) and DK752 and 125 RM R1 of each hybrid photosynthetically active and grain-filling period for all hybrids;
radiation (PAR) interception no effect on KN or kernel growth rate
by 1/3 with respect to intact
canopies

Borrás et al. (2009) 2005 4 hybrids and 4 inbreds – ?10 d before 50% anthesis Aiming at reducing – Defoliation increased the anthesis–
(Iowa) leaf area index (LAI) by silking interval and decreased the
?70%; accomplished by number of plants reaching silking and
removing all but the four plant growth rate around flowering up
uppermost leaves to 53%

Severini et al. 2007–2008 Iowa: popcorn public (R18, – 15 d before (Def−15) and Close to 100% (all but – KN: greater losses due to Def−15 (−13
(2011) (Iowa and IDS69, IDS91) and dent inbreds 17 d (Def+17) after 50% the top to −30%) than DF+17 (−5 to −19%);
Argentina) (B73, Mo17, N209) anthesis (beginning grain three or four leaves KW reduced the most by Def+17 (−9
filling) removed) to −38%)
Argentina: commercial hybrids
P625 and P802 (popcorn) and
AW190 (dent)

crop science, vol. 59, january–february 2019

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 14350653, 2019, 1, Downloaded from https://acsess.onlinelibrary.wiley.com/doi/10.2135/cropsci2018.04.0285 by Purdue University (West Lafayette), Wiley Online Library on [29/11/2022]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
more by GFD (up to −27%) than FD (up
Hanway, 1969; Vasilas and Seif, 1985a; Severini et al., 2011;

KN reduced most by 100% defoliation

100% defoliation at R2 (−23 to −48%);

Flowering defoliation reduced yield more (up KN reduced more by FD (up to −51%)
Yield component affected††

to −53%) than defoliation during grain filling than GFD (up to −14%); KW reduced

at V14 (−90 to −95%), followed by

KW reduced up to 71% by 100%

Tamagno et al., 2016; Battaglia et al., 2018), with KN
reductions as high as 62% with complete defoliation 10 d

defoliation at R2
after R1 (Egharevba et al., 1976), and 95% with complete
to −10%) defoliation at V14 (Battaglia et al., 2018). Although grain
yield effects in corn are mainly explained by changes in
the KN (Hall et al., 1981), grain yield reductions related
to a decline in KW with defoliation just before or at R2
(beginning of the grain-filling period) were widely docu-

§ Control treatment (not stated in the table) was included in all experiments. Vn, number of leaves completely unfolded; VT, anthesis; R1, silking; R2, blister stage; R3, milk stage; R5, dent stage.
mented in the past (Hanway, 1969; Egharevba et al., 1976;
defoliation at V14 and R2 reduced yield >85%
in most comparisons; 2013: hybrid ´ row ´
comparisons; 50% at V14 and 50% at R2

Tollenaar and Daynard, 1978; Echarte et al., 2006) without

and between 4 and 34% in 2013; 100%
reduced yield by 22% and 11% in 2012,
100% at V7 similar to control in most

changing the final KN. More recent studies showed that

partial and complete defoliation at R2 not only decreased
defoliationinteraction
Final yield#

KW, as expected, but also decreased KN (Tamagno et al.,

(up to −36%)

2016; Battaglia et al., 2018).

Echarte et al. (2006) tested plant defoliation treatments
to decrease assimilate availability during the grain-filling
period. These were established at 27 d after R1 to reduce
IPAR by 33%. Three to four leaves were left on each
plant after the defoliation treatment. For all hybrids tested,
defoliation did not affect KN per plant, nor the kernel
Defoliation timing§ Defoliation severity¶

100%: all timings, 50%:

growth rate relative to the control, but defoliation reduced

70 to 80% (upper 4
leaves always left
undefoliated)

mean final KW and grain-filling duration. Kernel weight

V14 and R2

was reduced by 38% for newer hybrids (released in 1993)

and 23% for older hybrids (released between 1965 and
1982). Echarte et al. (2006) concluded that a greater ear
demand relative to source capacity for newer hybrids
after anthesis (grain-filling
10–15 d before (flowering

explained their shorter effective grain-filling duration;

defoliation; FD) and 20 d

‡ Days to either 50% pollen shedding or physiological maturity. RM, relative maturity; GDD, growing degree days.
defoliation; GFD)

thus, newer hybrids had greater reductions in KW (lower

V7, V14, and R2

grain stability) in response to source reductions during

grain filling compared with older hybrids.
Eventually, 100% defoliation treatments at VT may
result in complete barrenness (Hanway, 1969). Test weight
(Dungan, 1934; Hicks et al., 1977; Lauer, 1994; Tamagno
113 and 120 RM

et al., 2016), shelling percentages (Hicks et al., 1977),

Maturity‡

ear weight (Hanway, 1969), and grain-filling duration

–

(Echarte et al., 2006) may also be negatively affected by

defoliation during the grain-filling period.
Table 2 provides a detailed summary of the data from
ACA2002, NT426, AX887, DK747
Hybrids or inbred lines†

different defoliations studies conducted in corn over the

P1360HR and P2088YHR

last 100 yr. Compiled information includes location, year

of study, hybrids and/or inbred lines used in the study
¶ Percentage of blades removed, if nothing else stated.

# As a function of the control (no defoliated) treatment.

and their respective RM classification, defoliation timing,

severity, and the impact of defoliation in final grain yield
and yield components.
†† KN, kernel number; KW, kernel weight.
† As quoted in the original manuscript.

HAIL DAMAGE EFFECTS ON THE

2012–2013
2012–2014
Study

DYNAMICS OF ANTHESIS, SILKING,

Table 2. Continued.

AND POLLEN SHEDDING

Hail at silking may bruise the silks, resulting in poor pollina-
(2016) (Argentina)

(2018) (Kentucky)
Tamagno et al.

Battaglia et al.

tion. Pollen shed and silking can be delayed by defoliation,

Reference

although the anthesis–silking interval (ASI) is less likely to

be changed ( Johnson, 1978). In experiments studying the

crop science, vol. 59, january–february 2019  www.crops.org 9


effect of early complete defoliation on tasseling–silking
synchronization, complete or near-complete defoliation at
or before V6 delayed the period from planting to anthesis
(Dungan and Gausman, 1951), silking (Lindstrom, 1935),
or both (Cloninger et al., 1974) by 2 to 8 d. Dungan and
Gausman (1951) reported maximum delays of up to 8 d in
pollen shedding with close-to-ground cuttings around the
V8 to V9 growth stage. Cloninger et al. (1974) indicated
that clipping plants delayed silking date by 8 d when plants
were clipped at or near ground level between V4 and V5,
compared with cutting at V3 and V6. In this case, the
maximum delay in silking was associated with the highest
grain yield reduction. Overall, cutting plants early (around
V3 for Cloninger et al., 1974; around V5 for Dungan and
Gausman, 1951) and rather severely resulted in maximum
delays in silking occurrence and minimum reductions in
grain yields and pollen shedding.
Fewer studies have addressed the potential impacts
that defoliations around the critical period in corn may
have on the dynamics of anthesis, silking, and ASI (Yao
et al., 1991). In a recent study conducted in Iowa, Borrás
et al. (2009) evaluated the impact of different treatments
aiming to alter plant growth around silking with the
objective to test the applicability of a silking simulation
model. A single defoliation treatment designed to reduce
LAI by ?70% was imposed to eight maize genotypes (i.e.,
four inbred and four commercial hybrids) around 10 d
before 50% anthesis. Defoliations were accomplished by
removing all leaves in the plant with the exception of the
upper four leaves, which were still expanding at that time.
Side-by-side undefoliated control treatments were also
established. Among other things, measurements collected
in this study included the anthesis and silking date, ASI,
plant growth rate (PGR, g plant−1 d−1) around silking,
LAI immediately after defoliation treatments allocation,
and IPAR (%) after maximum LAI (i.e., between 0 and
7 d post-50% anthesis). Defoliation treatments did not
affect the time to 50% anthesis, but genotypes differed
in their time to 50% silking and ASI response to defo-
liation. Compared with the control, defoliation increased
average ASI (range = 0.3–1.7 d, positive numbers indicate
anthesis followed by silking) in five out of seven compari-
sons, did not change the ASI in one comparison, and
changed the ASI from 0.3 to −1 d (i.e., silking before
anthesis). Moreover, the proportion of plants reaching
silking was also greatly reduced in the defoliated plots. In
fact, the remaining genotype was excluded from compari-
sons, as none of the defoliated plots reached 50% silking.
Compared with the control, and averaged across the eight
genotypes, defoliation treatments decreased LAI by 74%
(from 4.2 to 1.1 m 2 m−2), IPAR by 32% (from 86.5 to
59.0%), and PGR around silking by 37% (from 3.6 to 2.2
g plant−1 d−1). On a genotype-per-genotype basis, defolia-
tion reduced PGR between 21 and 53%. Clearly, IPAR
10 www.crops.org crop science, vol. 59, january–february 2019
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and PGR responses to defoliation were not proportional
to the total LAI removed by manual defoliations. In the
case of IPAR, light readings were taken at least 7 d after
defoliation treatments were imposed, with the remaining
leaves still expanding, and thus increasing their IPAR
readings, in between. Moreover, it is likely that the post-
defoliation photosynthetic efficiency of remaining leaves
was increased after defoliation (Dungan, 1934), thus
preventing the IPAR readings from decreasing in direct
proportion to the amount of area defoliated. Compen-
satory growth resulting from higher photosynthetic rates
of defoliated vs. undefoliated plants has been recognized
by others (Wareing et al., 1968; Briske and Richards,
1995; Collantes et al., 1998). Finally, and although sugar
levels in corn stalks are decreased by defoliation, a quick
and increased remobilization of stem reserves immedi-
ately after the defoliations, calculated to be ?19 to 24%
at anthesis (Kiniry et al., 1992), likely prevented PGR
around silking from declining at a greater rate.
HAIL DAMAGE EFFECTS AND PLANT
DISEASE PRESSURE
The direct impacts of hail on overall corn plant produc-
tivity can be increased indirectly by predisposing injured
plants to bacterial and fungal infection (Eldredge, 1935;
Robertson et al., 2011). Hail damage implications on corn
plant sanitary status have been studied for several diseases,
with the first studies being conducted in the United States
early in the last century. MacMillan (1918) estimated that
after a severe hail storm in Colorado, 19% of the affected
plants showed symptoms of small smut boils [Ustilago
maydis (DC.) Corda], whereas only 1% of the plants were
infected with the virus in an adjacent non-hail-affected
field. Dungan (1930, 1934) found that rotting of ears
after bruising by hailstones was worse at R3 (early milk
stage). Eldredge (1935) made similar observations, noting
that kernels damaged by hail had bruised or moldy spots
on the ear. These diseases may also increase mycotoxin
contamination, which represents a serious life-threatening
hazard both for livestock and humans (Robertson et al.,
2011). After intense damage caused by two major hail
storms at the R1 to R3 stage in 2009 in Iowa, Robertson
et al. (2011) collected corn ears within 48 h of combine
harvesting from 56 fields damaged by hail and 27 undam-
aged fields. Although levels of deoxynivalenol (DON),
zearalenone (ZEA), and fumonisins were detected in
grain samples obtained from both injured and uninjured
plants, mycotoxins were more frequently detected in hail-
damaged plants. Measured levels of DON and ZEA (2.63
and 0.43 mg kg−1, respectively) were 3.8 and 10.8 times
higher in kernels injured by hail, with no statistical differ-
ence in the level of fumonisins.
In the last 10 yr, fungicides have been promoted as
enhancers of plant tolerance to hail damage (Bradley and

Ames, 2010; Fernandez-Cornejo et al., 2014). Working
in Illinois with a 110-RM hybrid with moderate suscep-
tibility to grey leaf spot (Cercospora zeae-maydis Tehon &
E.Y. Daniels), Bradley and Ames (2010) studied the effect
of (i) mild defoliation severity (i.e., 25–30%) at the V12
stage on corn foliar diseases under natural disease pressure,
and (ii) two quinone outside inhibitor (QoI) foliar fungi-
cides (i.e., azoxystrobin and pyraclostrobin) applied at
VT on hail-damaged corn. Nontreated and nondamaged
controls were included in this experiment. Gray leaf spot
increased by 18% in defoliated plants in the first year, with
no differences in disease severity in the second year. Both
foliar fungicides reduced disease severity compared with
the nontreated control in the first year, but not in the
second. Application of two different QoI foliar fungicides
at VT to either defoliated or nondefoliated plants did not
improve yield compared with the nontreated control, a
lack of response that is in agreement with other studies
(Bradley, 2008; Conley et al., 2010; Sisson et al., 2016).
Bradley and Ames (2010) concluded by recommending
the use of a risk assessment criteria based on predictors
of disease, like the amount of corn stover at planting,
planting rate, hybrid RM, and resistance or tolerance to
key diseases like gray leaf spot, rather than potential hail
damage when considering foliar fungicide applications to
corn. Sisson et al. (2016), documented the effect of (i) hail
damage simulated with a string trimmer and ice machine
at VT and R2, and (ii) Headline AMP (pyraclostrobin +
metconazole) fungicide application timing (i.e., two levels:
“immediate” and “deferred,” applied ?3 and 8 d after the
defoliation event, respectively) on foliar disease severity
and grain yield. Common rust (Puccinia sorghi Schwein)
was the most important disease observed at most site years.
Sisson et al. (2016) reported lower foliar disease severity
at the R5 stage in hail-simulated plants at VT and R2,
compared with no hail damage in three out of the five site-
years, in contrast with the increased foliar disease severity
in defoliated plants reported by Bradley and Ames (2010).
In the other two sites-years, no significant differences in
disease severity between hail-damaged treatments were
observed. Finally, the combined effect of simulated hail
damage and fungicide application timing was evaluated.
Overall, across fungicides and site-years, simulated hail
damage at VT and R2 reduced yields by 25%. According
to Sisson et al. (2016), the lack of yield response to foliar
fungicide applications after simulated hail damage in the
critical period of corn yield determination may help to
decrease unjustified fungicide application.
SUMMARY AND FUTURE RESEARCH
DIRECTION
Corn response to hail damage mainly depends on defolia-
tion timing and severity. Damage to corn plants results from
reductions in corn plant stands and loss of photosynthetic
crop science, vol. 59, january–february 2019  www.crops.org 11
14350653, 2019, 1, Downloaded from https://acsess.onlinelibrary.wiley.com/doi/10.2135/cropsci2018.04.0285 by Purdue University (West Lafayette), Wiley Online Library on [29/11/2022]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
area. Older research suggested that leaves above and below
the ear were equally critical to grain yield, but recent
work with modern hybrids indicates that leaves above
the ear have a preponderant influence on the final grain
yield. Evidence of the plant productivity response to defo-
liation before the V10 growth stage is conflicting. If hail
damage occurs within 3 wk of emergence, a time when
the plant growing point is still below the ground, damage
has little to no effect on grain yield. However, removal
of 75 to 85% of the aboveground biomass within 3 wk of
emergence has reduced yields up to 33%. Yield response
to complete biomass removal between V2 and V6 can be
quite variable. Reported decreases between 5 and 44%
would seem typical, but this varies by hybrid type. For
example, complete defoliation at V3 to V4 increased grain
yield up to 56% in an early hybrid but reduced it by 7% in
a full-maturity hybrid. Once the growing point emerges
from the soil, around the V7 growth stage, complete defo-
liation can produce almost complete grain yield loss. Total
defoliation before V10 delays the period from planting to
anthesis, silking, or both by 2 to 8 d. Low-severity (e.g.,
25%) defoliation prior to VT (e.g., V12) may reduce grain
yield up to 30% but has little influence on corn yield at the
VT stage or later. More severe defoliation after the V10
to V12 stages gradually increases yield penalties to a peak
around the critical period, when complete defoliation can
cause complete or near-complete yield loss. After corn
passes into the reproductive phase, grain yield reductions
due to defoliation decrease progressively until maturity,
although losses can still be as high as 40% with complete
defoliations at the R3 to R4 stage.
The National Crop Insurance Service’s charts to
estimate the percentage of corn yield loss due to defo-
liation have undergone little change since the late 1960s
in the United States. Estimations in this system are based
on the “hail adjustor’s horizontal leaf method,” but this
method is less accurate and extended than the “collar leaf
method.” Adoption of the second method for all extension
and research activities in corn, included the assessment
of defoliation after hail damage, is warranted. Losses
resulting from corn defoliation due to hail damage prior
to V5 are calculated based on the need or lack of need to
replant corn stands, and not as a percentage of the final
grain yield loss. However, research conducted in the 1970s
in the US Corn Belt showed that defoliations between
V2 and V4 had the potential to not only reduce but also
to increase grain yield in some situations, something that
is not reflected in current adjusters’ loss charts. Most field
research assessing the impact of corn defoliations between
V5 and V9 includes the defoliation of the whole plant at
or near the soil level, a situation that is not comparable
with the information provided by adjusters’ charts, where
only the laminar part of the leaves has been subjected to
defoliation. When situations are comparable, differences

between adjusters’ charts and field research data are <15%.
Overall, a similar agreement can be found in most situ-
ations for the remaining part of the vegetative stage and
all the reproductive growth in corn. However, particular
comparisons, like the impact of complete defoliation
immediately before or after the occurrence of the VT to
R1 stage in corn inbreds lines or specialty crops like corn
with high oil content, seem not to be properly covered
by current adjusters’ charts. Moreover, at this time, the
validity of these charts to adequately describe situations
where interactions between two or more factors occurs is
somewhat dubious.
The two main yield components most attributed
to corn yield affected by defoliation are KN and KW.
Grain yield losses associated with defoliation around
VT to R1 were mainly explained by lower KN, with
reduction of up to 95% with complete defoliation near
the VT stage. Yield reduction by defoliation around R2
was related to a decline in KW in most studies, without
changing KN, likely reflecting decreased assimilate
availability during the grain-filling period due to lower
IPAR. However, recent research shows a reduction in
both the KW and KN with partial and complete defo-
liation at the R2 stage.
Defoliation by hail can increase plant predisposition to
bacterial and fungal infection. Increases in the frequency
of ear rotting, moldy spots, and small smut boils and in the
levels of DON and ZEA are examples of detrimental hail
effects on disease pressure and mycotoxin levels. Although
foliar fungicides have been promoted as enhancers of plant
tolerance to hail damage, their application around the
critical period does not appear to provide any benefit when
plant disease pressure is below the economic threshold.
There are limited data on the effect of management
practices like row width to corn yield response after
different defoliation timings and severities. Recent research
suggests that grain yield losses will be smaller with narrow
(38-cm) row spacing. Understanding differential genotypic
response is highly relevant. Several studies have shown that
corn maturity can play an important role in responses to
defoliation. Future studies on the effect of row width and
hybrid maturity on corn yield response after defoliation are
warranted to improve farm capacity to minimize the losses
to hail and other severe weather events.
Conflict of Interest
The authors declare that there is no conflict of interest.
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