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191
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2011

Corn Growth and Development


Lori J. Abendroth, Roger W. Elmore, Matthew J. Boyer, and Stephanie K. Marlay

IOWA STATE UNIVERSITY


University Extension PMR 1009 M arch 201 1
Authored by
Lori J. Abendroth,
Roger W. Elmore,
Matthew J. Boyer, and
Stephan ie K. Ma rl ay

Technical Support from


Sa rah A. Baune

Photography by
Iowa State University

Th is pub lication was


peer-reviewed by two
independent reviewers
using a double-bl ind
process.

Please refe rence


t his publicati on as:
Aben droth, L.J.,
R.W. Elmore, M.J. Boyer, Table of Contents
and S.K. Ma rl ay. 201 1.
Foreword .... ......1
Corn growth and
deve lopment. PM R 1009. Materials Used and Methods................. .. . ....... 1- 2
Iowa State University
Genetic and Management Advances . . . .. . .... .... ..3
Extension, Ames, Iowa .
Plant Development and Staging Methods ......4-7
Calculating and Predicting Corn Development... .............................. ................8- 9
... and justice for all
The U.S. Department of Agriculture Root Development .. .. ... ..10-11
(USDA) prohibits discrimination in
al l its programs and activities on the Seed Germination ............ .. .. ..... .. ...... 12
basis of race, color, national origin,
gender, religion, age, disability. political Vegetative Stages (VEto VT) . .. ........... . 13-27
beliefs, sexual orientation, and marital
or fam ily status. (Not all prohibited Reproductive Stages (R1 to R6) .. .. .28-41
bases apply to all programs.) Many
materials can be made available in Dry Matter Accumulation .. ...... .... .42
alternative formats for ADA clients.
To file a complaint of discrimination, Nutrient Accumulation ... ..43-46
write USDA, Office of Civil Rights,
Room 326-W, Whitten Building, Endnotes. . . . ... .. ... . ...47-49
14th and Independence Avenue,
SW, Wash ington, DC 20250-9410 or
call202-720-5964.

Issued in furtherance of Cooperative


Extension work, Acts of May 8 and
June 30, 1914 in cooperation with
the U.S. Department of Agriculture.
Gerald A. Miller, interim director,
Cooperative Extension Service,
Iowa State University of Science
and Technology, Ames, Iowa.
Foreword Materials Used and Methods
Corn Growth and Development replaces How a Corn Plant PLANT IMAGES
Develops, Special Report 48, by Iowa State University Plants used throughout this publication were grown in
Extension. 1 How a Corn Plant Develops was originally central Iowa. A commerc ial non-prolific hybrid, Golden
published in 1966 by Iowa State University agronomist Harvest 8529 CB/LL 7 was grown and photographed for all
John J. Hanway 2 and reprinted in 1971. Notably, this images. This hybrid conta in s transgenic traits integrated
publication included the first methodology 3 for stagi ng into its genetic background that provide resistance to
corn vegetative and reproductive development. Steven European Corn Borer and glufosinate herbicide. It has
W. Ritchie, John J. Hanway, and Garren 0. Benson rewrote a relative maturity (RM) of 108 days and falls w ithin the
How a Corn Plant Develops in 1982. The 1982 ve rsion recommended hybrid length for ce ntral Iowa. Growth,
conta ined a revised stag ing methodology, the Leaf Collar development, and appearance (such as leaf width and
method, whic h is now the most w idely recognized leaf angle) of this hybrid are similar to most dent hybrids
method in the United States and recommended for grown and marketed in the Corn Belt.8
practitioners 4 The dry matter and nutrient accumulation
figures in How a Corn Plant Develops were based on Seeds and plants photographed at vegetative (V) stages
research by Hanway.3. 5· 6 Since 1982, the publication VE, V1, V2, and V3 were greenhouse grown. Plants pho-
has had only one minor revision (1986) and numerous tographed at and after V6 were grown in pots outside to
reprints (1984, 1989, 1992, 1993, 1996, 1997, 2003, 2005, mimic environmental conditions typically experienced by
2008, and 20 10). Corn Growth and Development builds an Iowa corn crop. Multiple planting dates were used and
on the success of How a Corn Plant Develops and ranged from 23 April to 5 May 2009. Ears photographed at
includes who le plant photography, recent corn growth stages R1 to R6 were from field-grown (not potted) plants.
and development resea rch integrated into the text, and
'
new dry matter and nutrient accumulation figures. Photographed root systems were t aken from plants
grown in 5-gal (19 L) or 44-gal (166 L) pots to allow for
root excavation with minimal root breakage and loss that
often occurs with field e>«:avation. Plants in Figures 29, 30,
34, and 35 have less-than-expected root growth due to a
limited supply of plants grown in the 44-gal pots. A potting
soil comprised of 70 to 80% Canadian sphagnum peat
moss, perlite, dolomitic limeston e, gypsum, and a wetting
agent was used for the greenhouse pots (75% potting
so il: 25% sa nd mixture) and outside pots (1 00% potting
soil: 0% sa nd). The soil was compressed and watered
multiple times in each pot prior to seeding to reduce
so il settling. Seeds were planted approximately 1.5 to
2.0 inches (3 .8-5.1 em) deep. Plants were fertilized and
irrigated frequently to avoid any nutrient deficiency or
moisture stress.

Plants were staged often and leaves marked w ith paint


(not visible in photographs) to allow for leaf identifi cation.
Plants were tran sported to the Iowa State University
campus for studio photography with 3,000 images taken
over 14 studio sessions. Dissected plants are shown with
the leaves and ear shoots placed alternately (from side
to side) in succession as they natural ly appear on intact
corn plants.

Foreword and Materials Used and Methods


DRY MATTER AND NUTRIENT ACCUMULATION The primary (uppermost) ear shoot was cut at the base
A large-scale research project was conducted in Iowa of the shank once the shoot was visible above its respec-
during 2007 and 2008 investigating whether dry matter tive leaf sheath. Lower ear shoots rarely grew above their
(DM) and nutrient accumulation have changed in hybrids respective leaf sheaths and were therefore not separated
over the past 50 years? Data from two modern hybrids, from the stalk component. In add ition, lower ear shoots
Pioneer 10 33D11 (112 RM) and Pioneer 34A 15 (1 08 RM), did not produce grain; therefore grain data comes only
were compi led for the dry matter and nutrient accumula- from the primary ear. Plants were oven-dried at 140° F
tion figures (Figures 84-91) in this publication. Although (60° C) until weight loss ceased, then weighed. Dried plant
va riability in DM and nutrient accumulation existed tissue was ground for nutrient analysis with nitrogen (N),
among the two years and two hybrids, it was relatively phosphorus (P), and potassium (K) results published here
minor and the figures represent the expected response in elemental form .
of most hybrids and environments. The research project
was located on productive soi ls with an average grain Figures 84 to 91 were produced using Sigma Plot soft-
yield of 225 bushels/acre (bu/acre) (14.1 Mg/hectare [ha]); waren As shown, these figures have slight modifications
in comparison, the statewide yield average for each year from the orig inal research data 9 The sampling protocol
research was conducted was 171 bu/acre (1 0.7 Mg/ha) n did not include d issection of a few components at various
stages; therefore va lues were estimated as necessary. For
Hybrids were planted mid-May at 36,000 seeds/acre example, it was not feasible to separate the primary ear
(89,000 seeds/ha) to achieve a desired final stand of grain from the cob, shank, and husk fraction during R1
34,000 plants/acre (84,000 plants/ha). Phosphorus and and R2. Therefore, the grain portion of the graphic for R1
potassium were applied as needed in the fall or spring, and R2 is an approximation derived in part from published
and nitrogen was applied in the form of ammonium research on ear growth. 14• 15 The entire dataset for each
nitrate during the V3 to V4 developmental stages. Corn component was modeled using a polynomial smoothing
followed soybean each year; therefore, a recommended function in SigmaPiot in tandem with manua l adjustments.
nitrogen (N) ferti lizer rate of 150 lb N/acre (168 kg N/ha)
was applied. 12

Above-ground dry matter was collected and nutrient


content determined at 10 developmental stages during
each growing season . Whole plants were collected and
dissected at the following vegetative and reproductive
stages: V6, V9, V14, VT, R1, R2, R3, R4, RS, and R6. A total of
480 plants were dissected across all samp li ng stages. As
plants developed, the number of dissected components
increased to accommodate portions of the plant, such
as grain, that were not present early in the season. The
five plant components were: (1) stalk and leaf sheaths,
(2) leaf blades, (3) tassel, (4) ear shank, husk leaves, and
cob, and (5) grain. Leaf blades were cut at the leaf co llar,
leaving the leaf sheaths with the stalk component. Each
tassel was cut from the sta lk directly below the lowest
branch of the tassel.

2 Materials Used and Methods


Genetic and Management
Advances
Corn, as it is commonly referred to in the United States,
is known elsewhere in th e world as maize. The scientific
name for corn (maize) is Zea mays L. For more than two
decades, corn has been planted by U.S. producers on at
least 70 million acres (28 million hectares) with an upward
trend and occupies more acreage than any other grain
crop. 16 In terms of total world grain produced (weight
basis), corn lead s, followed by rice and wheat. 17

As with many plants, corn has been improved through


years of genetic and management research to meet the
ever-growing demands for food, feed, and alternative
uses. Advancements in corn management practices
and genetics have substantial ly increased grain yield
produced from an acre of land over the past 50 years. 18• 19
This increase in realized yields can be attributed in part
to genetic advancements that have increased stress
tolerance, improved resource capture, and increased
yield stabi lity. 18• 20 Genetic improvement has enabled
corn plants to be grown in close proximity, resulting in
greater yield potential from an acre of land 21 Contribut-
ing management factors made by the producer include
reduced tillage, earlier planting dates, increased plant understanding of crop growth and development equips
density per acre, herbicide use for timely weed control, practitioners to determine the status and health of
and synthetic fertilizers. the corn crop over the growing season and limit crop
stress when possible.
As plant breeders selected for high grain yield, intentional
and unintentional changes in plant architecture occurred, Growth and development are terms often used inter-
leading to improvements in grain production efficiency. 19 changeably, yet each has a distinct meaning. Growth
The modifications to hybrids can be documented when refers to the increase in size of an individual plant or
evaluated across decades. Today's hybrids differ primarily plant component. Development refers to the plant's
from those grown in previous decades by: ability to with- progression from earlier to later stages of maturity based
stand increased plant densities; more vertical leaf angle; on specific criteria that must exist to verify that the plant
reduced tassel size; near simultaneous occurrence of has reached a particular stage. Growth wil l vary signifi-
pollen-shed and silk emergence; increased plant stay- cantly due to temperature, moisture stress, weed pressure,
green; fewer barren ears; decreased protein and increased adequate fertility, etc. For example, two plants may each
sta rch content in grain; and altered root architecture that have five collared leaves and are thus developmentally
leads to deeper root systems.19· 20· 22 • 23 the same (VS); yet one plant may be shorter than the
other due to a stressfu l environment resulting in reduced
Producers and agronomists select hybrids for their specific growth. The progression of corn developmental stages
farms and optimize grain yields by employing science- is largely related to temperature and can be predicted.
based management practices that help protect the land Practitioners therefore more often discuss the status of a
w hile ensuring the highest output of grain. A thorough crop based on its development rather than growth.

Genetic and Management Advances 3


development. Once that is complete, its stage is based
Plant Development and on the development of the reproductive structures (ea rs).

Staging Methods Corn has male and female flowers separated by distance
on the plant as the tassel and ears respecti ve ly.
Corn expresses a determinate growth habit, which is
defined by the sing le sta lk terminating in the tassel, at top. Vegetative and reproductive developmental stages are
A determ inate plant differs from an indeterm inate plant determined on a w hole-field basis when 50% or more
in that vegetative structures (leaves and stalk) are initiated of the plants are at a particular stage. The Leaf Collar
prior to the initiation of reproductive structures (tassel method ' is used for stag ing veg etative (V) development
and ears). In contrast to corn, soybean varieties currently w hile reproductive (R) stages are based on established
grown in the centra l Un ited States are predominately inde- visual indicators of ke rnel development. See Table I.
terminate. Therefore, during most of th e growing season, Vegetative stages are designated w ith a "V" followed by
soybean plants are simu ltaneously staged accord ing to the tota l number of col lared leaves present. For example,
their vegetative and reproductive development, although a plant with one visible leaf collar is a V1 plant.
the reproductive stage is of most importance 24· 2s
The uppermost and final leaf on a plant va ries with
In corn, vegetative structures are initiated and then hybrid, planting date, and location, but most Corn Belt8
continue to grow wh ile the reproductive structures are hybrids produce 19 to 20 leaves. 27 The final leaf is simply
initiated and growing (Figure 1). Often, many portions of represented in Table 1 as Vn with n equa ling the number
the plant are growing but the plant is staged only by of the last leaf, such as V19 or V20. Two vegetative stages
what is identifiable at a specific point in time without do not require counting leaves: emergence (VE) and
dissection. The plant is first staged based on its vegetative tasse ling (VT). Reproductive stages are designated with
an "R" followed by the numbers 1 to 6.

Kernel ~ Growth

Kernel Initiation :

Ear Initiation and Growth

Tassel Initiation and Growth

:Internode Elongation :

Leaf Growth and Appea;ance



Leaf Initiation ....
-.
.. ..
c
Ia
Cll
Ia
c
....2::. V4 V6 V12 V16 VT/R1 R2 R3 R4 RS R6
~
"§ Cll
e'
Cll Cll
'-=' ....E
Fi~ure 1. Periods~f initiation and growth for individual above-ground plant structures from planting to physiological maturity (R6).16 Bold horizontal arrows represent the
pnmary penod of t1me when events occur; thin horizontal arrows represent possible variation. Kernel initiation refers to the initiation of florets, which may eventually become
kernels if pollinated and fertilized.

4 Plant Development and Staging Methods


Table 1. Corn vegetative and reproductive development stages. Slight to significant differences exist between the Leaf
Collar method and m ethods 1, 2, and 3, resulting in
Stage Common Name
occasional confusion among practitioners. Be aware of
VE Emergence the d ifferent stag ing systems, especia ll y w hen making
V1 First Leaf recommendations using a method other than the Leaf
"'> V2 Second Leaf Collar method. Convert between systems using values

-
·~
11:1

"'
~
C'l
V3
000
Third Leaf
000
shown in Table 2; BBCH conversions are not included
here but are published elsewhere 31 Re lative to the Leaf
Collar method, Horizontal Leaf wi ll be +0.5 to 2.5 stages
Vn nth Leaf
ahead and LeafTip w ill be +2.5 to 5.5 stages ahead. 32
VT Tasseling
R1 Silking Table 2. The Leaf Collar method compared to two other staging systems (Hori-
R2 Blister zontal Leaf and LeafTip) and plant height. 32 Plant heights can vary significantly
"'
>
'€ due to weather and management practices. Plant height is shown simply to serve
=
"1:1
e
R3 Milk as a reference to the three staging methods listed as well as allow for comparison
between methods of measuring height. Extended leaf height is measured from the
Ct. R4 Dough
"'
iCIC
R5 Dent soil surface to the tip of the uppermost leaf pulled straight, whereas canopy plant
height is measured to the natural bend of the uppermost leaves.
R6 Physiological Maturity

- •
lt:lHIJ :II If::

~i'J
Other stag ing methods exist in addition to the Leaf Collar
method: Leaf Horizontal Leaf Extended Canopy
(1) Horizontal Leaf also known as "Droopy" Leaf
Collar Leaf Tip Leaf Height Plant Height
• Used primarily by crop insurance adjustors. Inches
• Plant is staged based on the number of leaves V1 NA' 3.5 4 3
'
with 40 to 50% of their leaf area exposed along V2 3.0 5.0 6 4
w ith the tip of the uppermost leaf pointing V3 4.5 6.5 10 7
below horizontal 28
V4 5.5 0 8.0 15 10
(2) LeafTip 29 V5 6.5 21
9.0 14
• Used at times by the international scientific
V6 8.0 10.5 28 19
community.
V7 9.0 12.0 35 24
• Plant is staged based on the uppermost leaf tip
emerged from w ithin the whorl. V8 10.0 13.0 43 31

(3) BBCH 30 V9 11.5 14.5 52 39


• Used at times by the international scientific V10 12.5 15.5 60 47
community. V11 13.0 16.5 66 53
• Developed as a un iversa l sca le for stag ing V12 13.5 17.0 72 60
numerous crops. VB 14.0 17.5 78 66
(4) Plant height V14 15.0 18.0 84 73
• Used on pesticide labels often in specifying the NA**
V15 15.5 89 80
proper app lication timing of the product to
V16 NA' NA** 94 87
the crop.
V17 NA' NA** 99 93
• Leaf Co llar and Horizontal Leaf staging methods
may or may not be included on labels. V18 NA' NA** 103 100
V19 NA' NA" 106 103
V20 NA' NA" 107 105
*Not applica ble. The Horizontal leaf method is difficult to convertto the leaf Collar method atVl and
especially in later vegetative stages because of the vertical orientation of the upper leaves. The values for
Horizontal leaf method relative tothe leaf Collar method are different than published by the USDA-FCIC. 18
**Not applicable. The leafTip method is difficult to correlate to the leaf Collar method after V14."·"

Plant Development and Staging Methods 5


When using the Leaf Collar method, vegetative stages
are determined by counting the number of leaves w ith a
visible collar beginning with the plant's first leaf. A collared
leaf occurs when the leaf sheath (portion of leaf tissue
around the stalk) and leaf blade are visua lly distinct due to
the blade angling away from the stalk. Although a yel low
"line" wil l typically appear on the underside of a co llared
leaf (Figure 2), it is more reliable to stage based on the
upper side of the leaf where it opens away from the sta lk
(Figure 3). A leaf is fully collared when the leaf blade edges
are no longer crossed over one another around the stalk.
This methodology is similar to the classification of other
grass species, which in general is based on the appearance
of the collar or ligule (inner membrane present at the
junction of the leaf blade and sheath).

Corn grown at recommended Corn Belt8 plant densities


w ill have leaves placed alternately, each on the opposite
side of the stalk than the leaf directly below and above.
Figure 3. Collared leaves have slightlypale tissue at the point where the leaf opens
Collared leaves angle away from the plant at about a
away from the stalk.
45-degree angle. Leaf angle for the uppermost three to
four leaves is less, only about 30 degrees, as they remain
Counting collared leaves for determining the plant's
more ve rtical after becoming collared.
vegetative stage is possible until the lowermost leaves are
lost. Over t ime the lower leaves tear away, senesce, and
decompose due to stalk expansion, brace root formation,
or adverse weather (early season frost, hai l, flooding, etc.).
Once these lower leaves are no longer identifiable, a prac-
titioner must adjust with one of the following options:

(1) Cut or paint leaves earlier in the season to use as


an identification tool later. For example, leaf 5
can be cut or spray-painted when it first emerges
from the whorl. Later, when it is t he lowest leaf
remaining, a practitioner knows to start counting
at 5 and progress up the plant from there.

(2) "Split the stalk": Dig 1 to 2 representative plants per


field and split the stalk to identify to which stalk
node each of the remaining leaves are attached
(Figure 4). This is generally the most accurate
methodology to ensure the proper vegetative
stage is determined if option #1 is not possib le.

(3) Estimating the lowest vis ible leaf is the least


accurate approach but will often be correct
Figure 2. A"line"will exist on the back side of a collared leaf. given normal planting dates and environmental
conditions. The stalk node located flush with the
soil surface w ill typ ically be stalk node 6. Therefore,
the practitioner can identify leaf 6 by finding this
node and continuing to count nodes and their
respective leaves from that point upward.

6 Plant Development and Staging Methods


Options 2 and 3 are based on aligning collared leaves The last vegetative stage (VT) occurs when all branches
with their respective stalk nodes. For option 2, this is of the tassel are no longer enclosed by the upper leaves
done by splitting the sta lk lengthwise and examining but instead are fully visible and extended outward. Fifty
it. The key in this step is to identify the stalk node tissue years ago, the number of days separating pollen shed
apart from the internode tissue. An internode is the pithy (anthesis) and silk appearance was about a week, with
area between the stalk nodes and is light ye ll ow in co lor. silks emerging after the tassel began to shed. 19· 35· 36
Leaves 1 through 4 originate from stalk nodes located Although variability exists today among hybrids and
at the base of the stalk and when split, this area appears environments, most hybrids express the tassel and silks,
as a downward-facing triangle. Nodes 1 through 4 are and begin pollen shed at nearly, if not exactly, the same
tightly compressed and cannot be distinguished from time. 19 It is common today to have silks present and
one another, as no visib le internode tissue separates pollen shed occurring from the upper portion of the
them. Leaf 5 is attached to node 5 with approximately tassel before the lower tassel branches are fu lly extended .
0.1 to 0.3 inches (0.2 to 0.9 em) of internode tissue sepa- In the past, the tassel (VT) stage was defined as when all
rating it from node 4. 34 The amount of internode tissue tassel branches were fully extended with VT and silking
w ill vary between nodes 4 and 5 based on the planting (R 1) as separate and sequential occurrences. 1 This de-
date and when staging occurs. 34 Th e separation between scription and sequence no longer occur for most hybrids
nodes 4 and 5 becomes harder, if not impossible, to currently grown. In situations when the silks are out prior
distinguish later in the season . Therefore, the first visible to full extension of the tassel, the plant is defined as R1
internode tissue may be between nodes 5 and 6 instead despite the fact that VT has technically not occurred. 37
of between nodes 4 and 5. If the first visible internode
is greater than 0.4 inches (1.0 em) in length, it should be Reproductive stages are based on the grain development
assumed to be that between nodes 5 and 6 because that of the primary (uppermost) ear 1 and more specifically
length is not likely to occur between nodes 4 and 534 on the outer appearance and inner characteristics of
the kernels located in the middle of th e ear. Pulling back
the husk leaves attached to the ear shank allows for
examination of the kernels. Throughout the reproductive
stages, variation exists in kernel development from the
base (butt) to the tip of the ear. The base (butt) refers to
the portion of the ear closest to the shank and the tip as
the furthest (refer to Figure 56). The kernels located at the
base (butt) are more advanced developmentally com-
pared to those at the tip. Therefore, staging is based on
the kernels located in the middle of the ear as they serve
as an average for the ear. Although Corn Belt8 hybrids
typically produce only one ear per plant, it is possible to
have more than one ear on plants at field edges, at low
plant densities, or with prolific hybrids. If the plant has
more than one ear, the primary ear shou ld be used for
staging purposes.

Figure 4. Corn plant split lengthwise through the root system.

Plant Development and Staging Methods 7


Equation 2. Celsius growing degree day (GDDc)
Calculating and Predicting calculation for one day. Multiply the final GDDc va lue

Corn Development by 1.8 to convert to GDDF.


GDDc = [(TMIN + TMAx)/2]- 10
Corn development is correlated with the air temperature. TMIN = Minimum daily air temperature. If temperature
Therefore development varies from year to year if ca lendar
is less than 10° C, use 10 as TMIN·
days are only used to track progress. Yet development
becomes predictable within and across growing seasons TMAx = Maximum daily air temperature. If temperature
when evaluated using thermal time. The time required is greater than 30° C, use 30 as TMAX·
for corn to progress from one developmental stage to
another is based on the amount of heat accumu lated. Daily GDD va lues can be summed for a specific time
Thermal time represents the length of time the crop period and used to predict crop development. Some va ri-
spends within a defined temperature range considered ability, however, will exist between predicted and actual
optimum for that crop. development due to situations such as weather fronts
shifting the daily temperature significantly, prolonged
Several models for measuring corn thermal time exist38,39 cloudy periods, and severe shortages of water or nutrients.
although growing degree days (GDD) is the most used
and recommended for practitioners. Growing degree Throughout this publication, GDD val ues are calcu lated
days, modified growing degree days (MG DD), and grow- from VE (emergence) and not the planting date. Although
ing degree units (GDU) are terms used interchangeably the planting date is often more easily known than the
by practitioners for corn thermal time; GDD is used here. date of emergence, using it decreases the accuracy of
For corn, the range of air temperatures generally agreed the prediction in situations in wh ich air temperatures are
upon for growth begins at 50° F (1 ooC) and ends at 86° F optimum but other conditions are not. For example, a dry
(30° C) 40 although growth does occur outside these seedbed limits the capacity of the seed to germinate and
tempertures to varying degrees. The GDD calculation emerge, regardless if the temperature is appropriate. Many
assumes that corn development is consistent and linear resources, such as seed catalogs and other publications,
within the defined temperature range of 50° F and 86° F, start the accumulation of GDDs from the date of plant-
which is somewhat simplistic, but most practical. 38 These ing, which is something to be aware of when comparing
thresholds are used in calculating GDD as the minimum between this publication and others. Converting between
(TMIN) and maxi mum (TMAx) daily temperatures are set as resources is possible though, if seedbed and environ-
TMIN= 50° F (1 oo C) and TMAx = 86° F (30° C) 40 Equations mental conditions are favorable. A seedling will emerge
1 (° F) and 2 (° C), and the outer thresholds are the most approximately 90 to 120 GDDFfrom when it was planted 4 1
commonly used and recommended for corn. Corn can
accumulate between a minimum of 0 GDDFor a maxi- Leaf appearance can be predicted from VEto final leaf (Vn)
mum of 36 GDDF per day if temperatures remain :-::::50° F based on GDD accumulation (Figure 5). From VEto V1 O, a
or ;:::86° Fa ll day, respective ly. new col lared leaf appears approximately every 84 GDDF
accumulated. From V11 to Vn, each leaf appears more
Equation 1. Fahrenheit growing degree day (GDDF)
quickly at approximately every 56 GDDFaccumu lated 4 2
calculation for one day. Divide the final GDDFvalue by Once the plant is at R1 (si lking), its development can be
1.8 to convert to GDDc. predicted by using Figure 6.

GDDF= [(TMIN+ TMAx)/2]- 50


TMIN = Minimum dai ly air temperature. If temperature
is less than 50° F, use 50 as TMIN·
TMAX = Maximum daily air temperature. If temperature
is greater than 86° F, use 86 as TMAX·

8 Calculating and Predicting Corn Development


The relations hip between reproductive development and 1400
GD[) is more variable than that of vegetative development
and GDD. Within Figure 6, variability is most expected to 1200
occur after R4 w ith a range of +1- 50 GDDFper stage pos-
sible from that shown. 42 1000 ....... ·· ····· .. .

w
Reproductive development is determined based on the > 800

-
E
vis ual appearance of the outer and inner portions of the ...0
kernels (Table 1), yet these stages are only loosely linked c
LL
600 ....... ...... .
·~

to the actual changes taking place in grain moisture and c


(!)
dry matter. When either of these va riables (g ra in moisture .......
400 -~

or dry matter) are graphed re lative to accumu lated GDDs,


they progress in a linear fashion, w hereas the six R stages
200
(R1 - R6) do not. Therefore, practitioners must be aware
that the reproductive stages (R 1- R6) are not equa ll y
spaced by calendar days or GDDs (Figure 6).
E 2 4 6 8 10 12 14 16 18 20
Vegetative (V) Stage
Corn hybrids vary in the length of time necessary to reach <D
ll)
physiolog ica l maturity from emergence, w hether eva lu- ..t
ll)

ated by the number of calendar days or growing degree


Days after VE (Emergence)
days. Seed companies provide a framework of information
for practitioners to understand the length of time a specif- Figure 5. Vegetative development is pred ictable from VE (emerg ence)
ic hybrid requires from planting to phys iolog ica l maturity to Vn (final leaf) ba sed on growi ng degree day (GOD) accumulati on.41
The approximate number of cal endar days from VE to certain vegetative
(R6). The hybrid's re lative maturity (RM) is always provided,
stages is noted on the x-axis.43
and the number of GDDs needed from planting to si lking
(R1 ) and from planting to R6 are also often reported . The
re lative maturity is reflective of the length of t ime required 1400.-----~------~----~----~------.

for the hybrid to mature, but the RM value is not directly


corre lated to actua l ca lendar days; e.g., 120 RM does not 1200
mean 120 ca lendar days are required for maturity. Re lative
maturity values for hybrids grown in the Corn Belt8 are 1000
general ly between 90 and 120 RM. Hybrids w ith lower RMs .....
requ ire fewe r GDDs season-long to mature compared to D::
E 800
hybrids with higher RMs. .g
LL
cc 600
The total GDD req ui rement of a hybrid is divided approxi- (!)
mately in ha lf between vegetative and reproductive
400
deve lopment. 9· 44 Variation in development is related
to hybrid maturity, planting date, and environmenta l
200
conditions.45 A hybrid's re lative maturity (RM) does alter
the amount of t ime spent in vegetative and reproductive
deve lopment to some degree.
2 3 4 5 6
Reproductive (R) Stage

10-12 18-20 24-26 31 -33 64-66


Days after R1 (Silking)

Figure 6. Reproductive development is pred ictable from Rl (si lki ng) ba sed
on growing degree da y (GOD) accumulation. 41 The approximate number
of calendar days from sil ki ng (Rl ) to each reprodu ctive stage is noted on
th e x-axis. 43

Calmlating and Predicting Com Development 9


Seminal root system growth slows following emergence
Root Development of the coleoptile above the soil surface with maximum
size reached at approximately V2. The nodal root system is
The radicle is first to elongate from the seed followed by
visible at approximately V2 47• 48 and represents half of the
the coleoptile (shoot). Soon after the appearance of the
root mass byV3.1t becomes the dominant root system by
radicle and coleoptile, three to four additional roots
' V6 and for the remainder of the plant's life 48 The semina l
seminal roots, wil l elongate from the seed . Together,
roots are still distinguishable afterV6 and can serve as a
the radicle and seminal roots comprise the seminal root
diagnostic tool later in the season to identify the original
system, which helps establish the young seedling by
seeding depth, initial root health, or presence of compac-
absorbing water and nutrients from the soil.
tion zones.

Two distinct root systems exist in corn: semina l and nodal.


Nodal roots wi ll originate from each sta lk node below the
Noda l roots begin to develop at the co leoptilar node, the
soil surface and can be identified (Figure 8). Nodal roots
junction of the coleoptile and mesocotyl. Nodal roots are
that originate from stalk nodes above the soil surface are
consistently located 0.5 to 0.75 inches (1.3-1 .9 em) below
commonly referred to as "brace roots" since t hey ang le
the soil surface un less the seed was planted sha llow. The
downward to help brace the plant from w ind, although
placement of this root system is re lated to the perception
they function simi lar to other roots once they enter the
of incident light by the mesocotyl. 46 The length of the
soil. Brace roots may be visible by V9. The ir presence varies
mesocotyl varies therefore due to seeding depth and is
among plants and fields due to environmental conditions.
less for shallow planted and more for deeper planted corn
One to three sets of brace roots typical ly form on plants
(Figure 7). The recommended seeding depth for corn is
in production cornfie lds and are usua lly located at the
1.5 inches (3.8 em) or deeper to ensure the root system
nodes closest to the soil surface: nodes 6, 7, and 8 49 Node
develops properly and anchors the plant sufficiently.
6 brace roots are visible first and wi ll be fiush (para llel) to

Figure 7. ~eminal and no~al root development at different seeding depths: from 2.5 to 0.5 inches (6.2 to 1.3 em ), left to right respectively. Mesocotyl
leng~h vanes due to the different seedmg depths . Identifying the nodal root system apart from the seminal root system in the right-hand plant is not
possible because of shallow seeding .

10 Root Development
the soil surface while brace roots at nodes 7 and 8 will
ang le downward (Figure 9). Brace roots located at node 8
will typica lly not reach the soil surface. Approximate ly 70
nodal roots will originate from the stalk (below and above
ground) over the course of the season, beginning with the
first noda l roots to t he upper brace roots. 49

Root system formati on and arch itecture varies by genetics,


management practices, and environmental conditions. 48
Root s are distributed differently in t he so il profil e based on
thei r vertica l and horizontal distance away from the plant
and can be described in terms of their density (roots per
un it area) and length:

Density: In genera l, th e root density is greatest


closest to t he row and in the upper soi l profi le. 50· 51
During t he growing season, t he plant 's root density
increases until a maxim um is reached and t hen it
decreases. Th e developmenta l stage corre lated
w ith maximu m root density varies based on depth .
Density is maxim ized first in the upper profi le at Figure 8. Plant dissected lengthwise through the nodal root system .
approxi mately R1, w ith a gradual delay deeper in Each root originates from a specific stalk node. The seminal root system
is visible by the location of the depleted seed .
°
t he soil. 5 For exam pl e, maxi mu m root density does
not occur until approximately R3 w hen measured
3 feet (0.9 m) below the soil surface. 50
·.
Length: Root s elongate approximately 1 inch
(2.5 em) per day, w it h va ri ation among hybrids.47
At approximately V3, roots beg in entering t he
midd le of t he row (15 inches [38 em] from the
p lant). 50 The pl ant's root system reaches maxim um
depth bet wee n approximately R2 and R352 and
ca n extend more than 6 feet (1.8 meters) 53 deep
in soils th at do not have restrictions such as rock
layers or co m paction zo nes.

Figure 9. Brace root development at stalk nodes 6, 7, and 8.

Root Development 11
Seed Germination
Once planted, the seedbed needs to be suitable to ensure
good germination and eventual emergence including:
moisture, temperature, and seed-to-soil contact. Soils
should not be flooded or overly saturated with water
yet moist enough to allow for water absorption by the
seed. Water uptake occurs most rapidly through the
kernel tip and at a slower rate through the pericarp Figure 10. Whole and cross-sectioned kernels.
(seed coat); therefore, the embryo moistens first, followed
by the endosperm 5 4 A seed will absorb water until it is
The radicle, or primary root, elongates first from the seed,
approximately 30 to 35% moisture at germ ination when
followed by the coleoptile (shoot) growing in the opposite
it begins growth 54 Seeds exposed to prolonged coo l,
direction (Figure II). Under good conditions, elongation
wet cond itions germinate and develop slowly and wil l
of the coleoptile begins within a day of the emergence of
exhibit injury symptoms or die more frequently when
the radicle. The coleoptile grows approximately % inch up-
also subjected ·to soil-borne diseases,55 insect feeding, or
ward to the soil surface followed by differential mesocotyl
herbicides. To minimize these possibilities, corn should
growth. The mesocotyl is white internode tissue located
be planted when soil temperatures are near 50° F (1 C) oo between the seed and the coleoptilar node, and elongates
and rising. 55
to"push"the coleoptile to the soil surface. Mesocotyl elon-
gation continues until it perceives incident light near the
The seedbed must be managed to allow for good seed-
soi l surface 46
to-soil contact. Planter units open the soi l and place the
seed at the bottom of the furrow. Seed furrows should
be closed without creating a compaction zone or leaving
previous crop residue or soil clods next to the seed .

Specific seed components most often referenced are


the pericarp (seed coat), endosperm, and the embryo
(Figure 10). The aleurone, outer layer of the endosperm,
produces hydrolytic enzymes that activate when the seed
absorbs moisture from the surrounding soi l. Th e starch Figure 11. Corn germination with th e radicle and coleoptile elongating
and protein contained in the endosperm are broken from the seed. Seeds in this figure are colored pink due to a pesticide
down by the enzymes predom inantly into solubilized applied for protection from insect feeding and disease infection .
sugars and amino acids that are subsequently used by
the embryo. From the time of germination, the seedling
can survive for up to 14 days w ithout any additional
inputs apart from the endosperm. 48

12 Seed Germination
Vegetative Stages The number of calendar days between planting and emer-
gence varies and is primarily related to soil temperature,
moisture, and seed-to-soil contact. A seedling will emerge
VE: EMERGENCE though approximately 90 to 120 GDDF from the time it
was planted 41 Producers in the Corn Belts continue to
plant corn earlier, 56 often resulting in more exposure to
cooler temperatures and a greater number of calendar
days before the crop emerges.

The plant's first four to five leaves already existed within


the embryo and are visible with dissection 57 The co le-
optile tip opens at the soil surface, and these leaves are
able to grow upward through the co leopti le sheath that
had previously encased them. Photosynthesis begins and
is the process by which carbon dioxide and water are
converted into carbohydrates (such as sugars, ce ll ulose,
and starch) through numerous internal chemical reactions
dependent on energy from sun light.

The area of new cel l and structure initiation for corn is


referred to as t he growing point; it is botan ica lly termed
the apical meristem . The growing point is located at the
top of the sta lk and moves upward as lower internodes on
Figure 12. Emerged (VE) plant.
the sta lk become fully elongated. Add itional leaves (to the
previous four to five already present) are being initiated
A plant is defined as VE when the coleoptile emerges
now (Figure I) at the growing point. This is located at the
through the so il surface (Figure 12) until just prior to the
coleopti le base, 0.50 to 0.75 inches (1.3- 1.9 em) below the
first leaf collaring (Figure 13).The seminal root system
soi l surface.
wil l consist of the radicle (with branch roots) and
sem inal roots.

Figure 13. Corn seed ling development from germination to the second vegetative stage (V2) . Seeding depth is 1.5 inches (3.8 em) .

Vegetative Stages 13
V1: VEGETATIVE STAGE 1 V2: VEGETATIVE STAGE 2
Plants with their first leaf collared are defined as Vl (Fig- Plants with their first two leaves collared are defined as
ure 14). The tip of the first leaf is relatively oval shaped in V2 (Figure 15). Leaf initiation continues and formation of
contrast to all other leaves, which are pointed. This first leaf the nodal root system has begun. It is identifiable apart
serves as the starting point when counting leaves using from the seminal root system.
the Leaf Collar method. Leaves continue to be initiated
(Figure I) at the growing point, located below the surface.
The seminal root system is present and one or two nodal
roots may be visible, although none are vis ible in Figure 14.

Figure 14. Vl plant. Although three leaves are visible, the leaf collar is Figure 15. V2 plant. Although four leaves are visible, leaf collars are
distinguishable only on the lowest leaf. distinguishable only on the lower two leaves.

14 Vegetative Stages
V3: VEGETATIVE STAGE 3
Plants with their first three leaves collared are defined as V3
(Figure 16). The nodal and seminal root systems are about
the same size (length and dry matter) at this stage. Leaf
initiation continues at the growing point (Figure 1). Prior
to V3 and continu ing unti l V6, the plant is standing due
to the combined strength of leaf sheaths layered on top
of one another. The stalk remains below the surface at V3
although it is distinguishable with dissection (Figure 17).

Figure 16. V3 plant. Figure 17. Dissected V3 plant.

Figures 16 and 17. A V3 plant. A total of seven leaves are visible with dissection, although the upper leaves are within the whorl and not entirely vi sible
when the plant is intact.

Vegetative Stages 15
V6: VEGETATIVE STAGE 6
Plants with their first six leaves collared are defined as
V6 (Figure 18). The lower leaves are more weathered
and become increasingly harder to identify and count as
they tear away from the expanding sta lk and decompose.
All leaves are initiated by V6 although many are too sma ll
to see without magnification. Each leaf originates from a
stalk node with internode tissue separating the nodes.
A minor amount of internode elongation began prior to
V6 with the majority occurring from this point forward
(Figure 1). The growing point has now transitioned from
below to above the soil surface due to internode elonga-
tion (Figure 19). The nodal root system is dominant now
with the root mass approximately one third of the plant's
tota l biomass 47

Ear shoots, one of which will develop into a harvestable


ear, are being initiated and growing along the sta lk at
various nodes (Figure I). Ear shoots are first present at
lower stalk nodes as these are initiated first with upper
ear shoots following. For example, in Figure 19 the ear
shoot located at node 5 is larger than those located at
nodes 6 or 7 because it was in itiated ea rl ier and has had
more t ime to grow.

Although the primary ear shoot is not yet visible, it is


initiated at approximatelyV6; 58 magnification will be
necessary to see it. The primary ear is typ ically located
at nodes 12, 13, or 14.59 The potential size of an ear is a
function of the number of kernel rows around the ear Figure 18. V6 plant.
and the number of kernels per row. The row number
wi ll be even (versus odd) because initia l rows divide Figures 18 and 19. A V6 plant with the lowest two leaves lying on the
lateral ly, form ing two rows each. Row number is deter- soil surface due to stalk expansion . A total of 161eaves are vi sible with
mined shortly after the ear is initiated, approximately V7 58 dissection, yet the upper leaves are within the whorl and not visible
when the plant is intact. The leaf area not exposed to the sun is pale
Most hybrids grown commercially have 16 or 18 kernel
ye llow. Three ear shoots are visible on this plant, with the lowest ear
rows per ear. The row number is strong ly related to a
shoot located at node 5.
hybrid's genetics and impacted only by serious environ-
mental factors such as drought, nutrient deficiencies,
and improper herbicide applications. (R1 ), 6tl which is approximately V15 orV16. The total
number of potentia l kernels per ear is related to growing
The initiati on of flo rets ("potent ial kerne ls") w ith in a row conditiDns prior to silking (R1) whi le the number of
does not occur sim ultaneously; potential kernels near harvestable kernels is related to the conditions during
the base (butt) are first and the t ip potential kerne ls last.60 and after R1.
A floret describes a female flower prior to ferti lization
although it is simply referred to here as a "potential The tassel is initiated at approximatelyV6 (Figure 1),
kernel:' Potential kernels will begin to be initiated now although it will not be visible apart from magnification.
and continue until one week prior to silk emergence It is identifiable w ith plant dissection byV7 (Figure 20).

16 Vegetative Stages
Figure 19. Dissected V6 plant.

Figure 20. Upper portion of the stalk, with the tasse l visib le,
from a V7 plant.

Vegetative Stages 17
V9: VEGETATIVE STAGE 9
Plants with their first nine leaves collared are defined as V9
(Figure 21 ). Therefore, approximately half of the total leaves
on a plant are now collared assuming it will have 19 to
20 total. 27 The lower two to three leaves are usually fully
or partially decomposed now due to being torn from stalk
expansion and brace root formation. Without leaf 1 visib le
and serving as a starting point in the Leaf Collar method,
an additional step is needed to identify the remaining col-
lared leaves. Determining the vegetative stage is possible
by previously marking the leaves, splitting the stalk, or by
estimation; refer to page 6.

The growing point continues to move upward as lower


internodes become fu lly elongated (Figure 22). Brace roots
may be present at the soil surface and begin to help an-
chor the plant and obtain water and nutrients. The tassel,
visible with dissection, is pale yellow and rapidly growing
(Figure 23).

All ear shoots are initiated now and actively growing (Fig-
ure 24). Ear shoots will vary in size relative to one another
due to when they were in itiated and the ir location on the
stalk. Although the lowermost ear shoots were formed
first, their growth does not continue once the uppermost
ear is established as that becomes dominant over those
below. An ear shoot can be found at each above-ground
stalk node except for the upper six to eight nodes in Corn
Belt8 hybrids. 59

Figure 21. V9 plant.

Figures 21 and 22. AV9 plant with nine leaves collared although leaves
1 to 3 are no longer present on thi s plant. Atotal of 20 leaves are vi si ble
with dissection, yet the upper leaves are within the whorl and are not
vi si ble when the plant is intact. Th e leaf area not exposed to the sun is
pale ye low. Eight ear shoots are present, with the primary (uppermost)
ear shoot located at node 13. Th e lowest ear shoot is at node 6; the ear
shoot ~ ocated at node 5 (Figure 19) is not identifiable on this plant, likely
due to brace root formation .

18 Vegetative Stages
Figure 22. Dissected V9 plant. Figure 23. Upper portion of the stalk,
with the tassel visible, from a V9 plant.

Figure 24. Ear shoots from a V9 plant.


Shoots are arranged from the lower to
upper stalk nodes (left to right); nodes
6 to 13 respectively. The ear shoot at
node 13 on this plant will grow rapidly
and become the primary ear. The ear
shoot present at node five in Figure 19
was not present or too small to locate
on this plant.

Vegetative Stages 19
V12: VEGETATIVE STAGE 12
Plants with the 12th leaf co llared are defined as V12
(Figure 25). Approximately 10% of a plant's total dry matter
is now accumu lated (Figure 85).

The lower th ree to fou r leaves w ill not be present due to


sta lk expansion and subsequent decomposition. As w ith
earlier stages, w ithout leaf 1 serving as a starting point in
the Leaf Collar method, an additional step is needed to
identify the rema ining col lared leaves, refer to page 6.

The growing point continues to move upward as lower


internodes become ful ly elongated (Figure 26). The tassel
is rap idly growing and becoming more yellow-g reen in
co lor (Figure 27). The uppermost ear shoots are growing
much faster than the lower shoots (Figure 28).

Figure 25. V12 plant.

Figures 25 and 26. Twelve collared leaves exist although leaves 1 to 4


are no longer present on this plant. Twenty-one leaves are present with
dissection, yet the upper leaves are within the whorl and are not visible
when the plant is intact. The leaf area not exposed to the sun is pale
yellow. Eight ear shoots are present, with the lowest ear shoot at node 7.
The ptimary ear shoot is at node 14 on this plant whereas in Figure 22 it
was node 13; this is simply due to plant variation. The ear shoots located
at node 5 (Figure 79) and node 6 (Figure 22) are not identifiable on this
plant, likely due to brace root formation .

20 Vegetative Stages
Figure 26. Dissected V12 plant. Figure 27. Tassel from a V12 plant.

Figure 28. Ear shoots from a V12 plant;


nodes 7 to 14.

Vegetative Stages 21
V15: VEGETATIVE STAGE 15
Plants with the 15th leaf collared are defined as V15
(Figure 29) . Approximately 25% of a plant's total dry matter
is now accumu lated (Figure 85). The lower three to four
leaves wi ll not be present due to decomposition. Th e
growing point continues t o move upward as the lower
internodes elongate (Figure 30). The tassel continues to
grow (Figure 31 ).

The upper ear shoots continue to grow whi le growth


of the lower ear shoots has all but ceased (Figure 32).
The upper two ear shoots are now simi lar in size, but the
primary ear wil l receive more resources and dominate.
The number of potential kernels (florets) per row is at or
near fin al now as kernel initiation is finished one week
prior to si lk emergence (Rl )60 Since most Corn Belt8
hybrids have 19 to 20 totalleaves, 27 this equates to
approximately V15 or V16. A floret describes a female
flower prior to ferti lization although it is simply referred
to here as a "potential kernel :'

The total number of potential kerne ls per ear is re lated to


g rowi ng con ditions prior to silking (Rl ), wh ile t he number
of harvest able kernels is in response to cond itions during
and after Rl . The ear will have 700 to 1,000 potential
kernels, 61 w hich develop into kerne ls once fertilized.
Typically, 450 to 550 kernels per ear are expected at har-
vest, based on recommended practices and a favorable
environment. 62 Genera lly, more gra in yie ld is produced
per acre w ith a rel atively hig h plant density combined
w ith an adequate number of kernels per ear rather than
a low plant density and a high number of kernels per ear. Figure 29. V15 plant.
Th erefore, the number of kerne ls per ear harvested shou ld
be less than the existi ng potential (700- 1,000) to maxi- Figures 29 and 30. Fifteen collared leaves exist although leaves 1 to 3 are
m ize total gra in yie ld per acre. no longer present on this plant. Twenty leaves are visible with dissection;
the upper leaves are within the whorl and not entirely visible when the
plant is intact. The leaf area not exposed to the sun is pale yellow. Eight
The silks of the uppermost ears are now starting to elon-
ear shoots are present on this plant, with the upper two significantly
gate from near the base (butt) potential kernels (Figure 33).
larger than those below. The lowest ear shoot is at node 6; the ear shoot
Sim ilar to kernel initiation, not all si lks grow simultaneously at node 5 (Figure 19) is not identifiable on this plant likely due to brace
with si lks attached to t he base kernels first and tip kernel root formation. The primary ear shoot is at node 13 on this plant whereas
sil ks last 60 in Figure 21i it was node 14; this is simply due to plant variation. Root
growth, as shown, is less than expected at this developmental stage
and is attributed to space limitation s for this specific plant (refer to the
Materials Used and Methods section).

22 Vegetative Stages
Figure30. Dissected V15 plant. Figure 31. Tassel from aV15 plant.

~
00
0 ...... 0
0
....
0::
0
-o }-rip
a- 0
Kernels
....
~

0 00
0

;::::
0
"' ......
0
-Silks
c:
0
.... a-
0

I' "' "'


}-··"
00 0
0

.. ....
f
I rSh•nk
a- 0 Kernels
0

.... "' ""'


0

l
0 '

I \
"'
0

' '
"'
0
I!
0 0 ll i 0

mm inch 0 0

mm inch
Stalk Node: 6 7 8 9 10 11 12 13

Figure 32. Ear shoots from a V15 plant; nodes 6 to 13. Figure 33. Three uppermost ears, from nodes 11 , 12, and 13. The primary ear
is on the right.

Vegetative Stages 23
V18: VEGETATIVE STAGE 18
Plants with the 18th leaf collared are defined as Vl8
(Figure 34). Approximately 35% of a plant's total dry
matter is now accumulated (Figure 85). The lower four
to five leaves wi ll not be present due to decomposition.
The upper leaves remain more vertical, at an approx-
imate 30-degree angle, compared to t he lower leaves
at approximately 45-degrees. Nearly all internodes are
fully elongated except for those on the uppermost
portion of the sta lk (Figure 35). The tassel continues to
grow and is nearly fu ll size (Figure36andFigure41).

The upper two ear shoots are sim ilar in size (Figure 37).
Wit h t he remova l of the husk leaves, it is clear that si lk
elongation is most progressed on the primary ear
(Figure 38), although the two uppermost ears are fairly
sim ilar in size (Figure 39). Sim il ar to kerne l initiati on, not
all silks grow simu ltaneous ly, w it h silks attached near
the base (butt) kernels beginning first and tip kernel
si lks last.60 Sil ks are visible from both base and t ip
kerne ls (Figure 38), although the t ip kernel si lks are
shorter due to their delayed start. Most often, the
pri mary ear is located at node 13 but it ca n easil y be
greater or less by one node due to hybrid, planting
date, and environmental differences. 59

Figure 34. V18 plant.

Figures 34 and 35. Eighteen collared lea ves exist although leaves 1 to 5
are no longer present on this plant. Twenty-one leaves are visible with
dissection . Six ear shoots are present on thi s plant, with th e upp er two
significantly larger than the othe rs. The primary ear shoot is at nod e 13 on
this plant and is visible above the leaf sheath . The lowest distinguishable
ear shoot is at node 8 with lowe r ea r sho ots not identifiable on thi s plant,
likely due to brace root formation . Root growth , as sho wn, is less than
expe!ted at this developmental stag e and is attributed to spa ce lim itations
for this specific plant (refer to the Materials Used and Methods section).

24 Vegetative Stages
Figure 35. Dissected V18 plant. Figure 36. Tassel at V18 .

Figures 38
and 39. Three
uppermost ears,
from nodes11,
12, and 13.

Figure 37. Ear shoots from a V18 plant; nodes 8 to 13. Figure 38. Three uppermost ears with Figure 39. Three uppermost ears
silks, primary ear on right. without silks, primary ear on right.

Vegetative Stages 25
VT: VEGETATIVE STAGE TASSEL
Plants with all branches of the tassel fully visible, extended outward, and not held in by
the upper leaves, are defined as VT (Figures 40 and 41 ). A plant is defined as VT regardless
of whether it has begun shedding pol len (anthesis) or not (see Figures 42 and 43) as it is
based solely on whether or not the tassel is completely visible.

Plants atVT have Vn leaves (n= final leaf) and are at maximum or near-maximum height.
Most hybrids grown in the Corn Belt8 will have a total of 19 to 20 leaves prior to thetas-
sel 27 The tassel is at maximum size (Figure 41) and dry matter.

Although the tassel is an easy structure to identify for staging purposes, the occurrence
of pol len shed (anthesis) is more important to document. The shedding of pollen is a
determin ing factor in whether or not silks become poll inated and potentia l kernels fertil-
ized. All branches of the tassel may not be fu lly extended above the upper leaves before
the anthers on the main branch start shedd ing pollen (Figure 44). Also, si lks will often be
visible before the tassel is fu lly extended above the upper leaves; if this occurs the plant
should be defined as Rl despite VT not techn ica lly occurring first (refer to page 7).3 7

The length of the pollination window differs based on whether it is for the whole fie ld
or for an individual plant. Plants with in a field do not all beg in or end pollen shed at the
same time due to plant variabi lity. Most fields wi ll have pol len shed occurring for seven
or more days 61 However, the greatest production of pollen from that field exists for a
shorter time period of approximately four days.61 An ind ividua l plant at peak pollen pro-
duction can re lease one-ha lf a million or more pollen grains per day, although variation
exists among hybrids and plant densities 61 Figure40. Tassel from a VT plant.

V2 V3 V6 V9 V12

Figure 45. Vegetative development from V2 to R6. Maximum plant height is reached at approximately R2 .

26 Vegetative Stages
Figure 42. Tassel at VT, prior to pollen shed.

Figure44.
Main stem of the
tassel. Anthers
are exserted and
Figure 43. Tassel at VT during pollen shed. Anthers shedding pollen,
are exserted from the flowers and releasing pollen although the
Figure41. Tassel growth beginning at V7 and complete at VT . The grains (not visible) . Tassel is angled to the side to pollen grains are
internode below the tassel elongates as the tassel grows. see the anthers in more detail. not visible here.

Vegetative Stages 27
Reproductive Stages Rl
I
R2
I
R3 R4
I
RS R6
I

Reproductive development is based on


examination of kerne ls from the middle of I }Emb<yo

• ' ' ''


Side

••
the primary ear. Kerne ls change in several
significant ways following fertilization:
change in color from white to deep yellow, I} "
Non-

' ''
Embryo
decrease in moisture content, development Side

of the embryo, and increased starch
accumulation. These changes are all Figure 46. Whole kernel development. The embryo and non -embryo sides of each kernel
visible in Figures 46 to 49. are shown once they are distinguishable.

O
• '
,1 III '~
== Endosperm
Emb<yo
Black Layer

Figure47. Longitudinal cross-section of kernels.

Figures 46, 47, and 48. Whole and cross-sectioned


kernels from Rl to R6. Two images of R3 and RS are
shown due to the range possible based on time of
sampling within each stage. The embryo is visible in
the cross-section images as yellow-white tissue in the
lower right side of each kernel (Figure 47) or the center
(Figure 48) . Starch accumulation increa ses over time
and the kernel interior becomes white at the top
progressing down toward the base, or tip. Figure 48. Planar cross-section of kernels .

Figure 49. Cross-section of primary ears from Rl to R6 . Two ima ges of R3 are shown due to the range possible based on time of sampling within this stage.
The embryo and non -embryo sides of each ear are shown once they are distinguishable. The cob pith is always white although the outer perimeter of the cob
varies among hybridsfrom white to red; here it is pink.

28 Reproductive Stages
R1: REPRODUCTIVE STAGE 1 (SILKING) Although kernels are pol linated and fertilized during R1,
Plants defined as R1 must have one or more silks the ear is at the beginning of a rapid elongation period
extending outside the husk leaves (Figure 50). Plants are °
and is only 40 to 45% of its finallength. 14• 6 Kernels at
at maximum or near maximum height (Figures 45 and 51) R1 are nearly encased in glumes (termed botanically as
and have near maximum vegetative dry matter (Figure 85). sepals). When viewed from the side, the kernels appear
Determining the reproductive stage of the crop at and pointed because of the silk scar, which is where the si lk
after R1 is based solely on the development of the primary was attached. The outside of the kernel is white and the
ear. Silking (R1) is the only reproductive stage defined not inside clear, due to its high water content (Figure 53).
,,
on the characteristics of individual kernels but rather on The embryo begins to form fo llowing ferti lization, yet
I
the appearance of silks outside the husk leaves. it is not distinguishable without magnification. As the
plant approaches R2, kernels expand and have angled
The si lking period is the most sensitive period for the sides and a flatter top.
crop; stress at this time can reduce kerne l number per
ear63 Silks on the primary ear must be present while pol-
len shed (anthesis) occurs for successful pollination and
ferti lization. Synchronization between pollen shed and
si lking is important for obtaining high gra in yields. 63

During R1, both pollination and fertilization occur.


Pollination refers to the transfer of pollen gra ins (male
reproductive cell) to the silks that are each individual ly
attached to an ovary (female reproductive cel l). Fertiliza-
tion is the joining of these two reproductive cells to
create an embryo.

Each si lk is attached to one potential kernel. A poller1


grain can land anywhere on an exposed silk and may
germinate leading to fertilization. Silks remain receptive
to pol len for a min imum of five days after they emerge. 64
The first si lks to emerge from the husk leaves are those
attached to potential kerne ls near the base (butt) of the
ear. Si lks attached to potential kerne ls at the ear tip
are last to emerge and may not be pollinated if pol len
shed has ended. Some potential kernels w ill simply
not develop into harvestable kernels due to a failure in
pollination or fertilization; these kernels wil l be vis ible
on the ear as smal l, undeveloped white mounds.

Figure SO. Primary ear at Rl shown with and without husk leaves and
silks. The base of the shank is the point of attachment to the stalk.

Reproductive Stages 29
Figure 51. R1 plant. Figure 52. R1 plant dissected.

Figure 51 and 52. R1 plant with silks extending from the primary ear. This plant has 21 leaves, although the lower seven are no longer present. The
primary ear shoot is at node 13 on this plant and the lowest distinguishable ear shoot 1s at node 8. Ear shoots below node 8 are not identifiable, likely
due to brace root formation .

30 Reproductive Stages
Figure 53. Kernels from Rl plant. Kernels are arranged L toR : whole,
planar cross-section, and longitudinal cross-section.

Figure 54. Three uppermost ear shoots from Figure 55. Three uppermost ears with silks; Figure 56. Three uppermost ears without silks;
Rl plant; primary ear on right. primary ear on right. primary ear on right.

Figures 54, 55, and 56. Three uppermost ear shoots and ears from Rl plant. Staging is based solely on the primary ear. A slight change in silk color of the
primary ear silks marks the transition area from inside to outside the husk leaves for each silk.

Reproductive Stages 31
R2: REPRODUCTIVE STAGE 2 (BLISTER) Kernel abortion occurs primarily during R2 and R3 and is
related to an inadequate carbohydrate supply from the
plant. The kernels fertilized last are those aborted first
resu lting in the tip kernels most often aborted.

Silks outside the hu sk leaves are drying and chang ing in


color from tan to light brown (Figure 59). Silks wi ll naturally
detach from their respective kernels following fertilization
Figure 57. Kernels from R2 plant. Kernels are arranged L toR : whole,
and can be seen if the husk leaves are removed and the
planar cross-section, and longitudinal cross-section .
ear shaken.

Figure 58. Glumes partially surrounding each kernel are visible when
looking at an ear cross-section .

R2 occurs approximately 10 to 12 days after R1 (Figure 6).


Plants defined as R2 have kernels that look simi lar to
"bl isters." Kernel growth begins fol lowing fertilization and
consists of a rap id increase in water content with 85%
kernel moisture at the beginning of R2 65 Gra in dry mat-
ter accumu lation is minimal now (Figure 84). Plants at R2
have reached maximum height (Figure45) and maximum
vegetative dry matter (Figure 85). The ear is now at its final
(maximum) length.14

The g lumes su rround ing each kernel are visual ly less


prominent now as the kernels expand beyond them
Figur 59. Primary ear at R2 shown with and without husk leaves and
(Figure 58). Kerne l expansion is occurring with the kerne ls silks. The base of the shank is the point of attachment to the stalk.
now rounded although space still exists between the ker-
nel rows. The si lk scar on the kerne l top is still identifiable
as a circle. The outside of the kernel is ivory colored, and
the inside remains a clear liquid (Figure 57) . The embryo is
growing, yet is not distinguishable without magnification.

32 Reproductive Stages
Figure 60. Three uppermost ear shoots; primary Figure 61. Three uppermost ears with silks; primary ear Figure 62. Three uppermost ears without
ear on right. on right. silks; primary ear on right.

Figures 60, 61 and 62. Three uppermost ear shoots and ears from R2 plant. Staging is based solely on the primary ear. The change in silk color to brown
marks the transition area from inside to outside the husk leaves for each silk.

Reproductive Stages 33
R3: REPRODUCTIVE STAGE 3 (MILK)

Figure 63. Kernels from R3 plant. Kernels are arranged L toR : whole,
planar cross-section, and longitudinal cross-section .

R3 occurs approximately 18 to 20 days after Rl (Figure 6).


Plants defined as R3 have kernels with a "m ilky" interior
and explode easily when pressure is applied. Kernel
moisture is approximately 80% at the beginning of R3. 65
Th e outside of the kernel is yellow and the inside is white
and somewhat translucent. The glumes, which previously
encased the kernels, and the si lk scar, are barely visible.
Kernels completely fill the space now between kernel
rows. Starch accumu lation is increasing resulting in greater
kernel dry matter (Figure 84). The embryo and endosperm
are now distinguishable (Figure 63) . Sim ilar to R2, kernel
abortion can still occur if carbohydrate supply from the
plant is inadequate during R3.
Figure 64. Primary ear at R3 shown with and without husk leaves and
silks. The base of the shank is the point of attachment to the stalk.

34 Reproductive Stages
Figure 65. Three uppermost ear shoots; primary ear Figure 66. Three uppermost ears with silks; primary ear Figure 67. Three uppermost ears without silks;
on right. on right. primary ear on right.

Figures 65, 66, and 67. Three uppermost ear shoots and ears from R3 plant. Staging is based solely on the primary ear; lower ears are not developing further. The change
in silk color to brown marks the transition area from inside to outside the husk leaves for each silk.

Reproductive Stages 35
R4: REPRODUCTIVE STAGE 4 (DOUGH)

t t t I
Figure 68. Kernels from R4 plant. Kernels are arranged L to R: whole on
embryo side, whole on non -embryo side, planar cross-section, and longi-
tud inal cross-section .

R4 occurs approximately 24 to 26 days after Rl (Figure 6).


The plant is defined as R4 when the consistency of the
kernel interior is similar to "dough:' The outside of the
kernel is deep yel low and the inside is white and less
translucent than at R3 (Figure 68). Kernels have a matte
finish (compared to their previous glossy appearance),
the tops are flattening, and more pressure is needed
to explode compared to R3 kernels. Kerne l moisture is
approximately 70% at the beginning of R4. 65 Near t he
end of R4, kerne ls (often those near the base) beg in to
indent at their top due to increasing starch deposition
and moisture loss. The cob co lor is hybrid specific and
can remain wh ite (as it was prior to R4) or change to
pink or red (Figure49). Ears have husk leaves that are
beginn ing to turn brown on the edges. Figure 69. Primary ear at R4 shown with and without husk leaves and
silks. The base of the shank is the point of attachment to the stalk.
Starch accumulation continues to increase resulting in
greater kerne l dry matter (Figure 84). Stress during this
stage will not resu lt in aborted kerne ls, but instead a
reduction in kernel weight because less starch is accu-
mulated. An environment that is not stressful for plant
development wi ll result in increased carbohydrate
(starch) accumulation and heavier kernels than a
stressful environment.

36 Reproductive Stages
Figure 70. Three uppermost ear shoots from R4 Figure 71. Three uppermost ears with silks; primary Figure 72. Three uppermost earswithout silks;
plant; primary ear on right. ear on right. primary ear on right.

Figures 70, 71, and 72. Three uppermost ear shoots and ears from R4 plant. Staging is based solely on the primary ear; lower ears are not developing further.
The change in silk color to brown marks the transition area from inside to outside the husk leaves for each silk.

Reproductive Stages 37
RS: REPRODUCTIVE STAGE 5 (DENT) Table 3. Progression of milk line during RSwith approximate percent moisture,
dry matter, growing degree day (GDDF) and days for each substage. 70 Grain moisture
and dry matter (DM) values are an average and variation of at least +/-2%is
expected with all except for DM at R6, which is always100%. Growing degree day
and calendar day valuesare from Figure 6.43

I ' I I
Figure 73. Kernels from RS plant. Kernels are arranged L toR : whole on
embryo side, whole on non -embryo side, planar cross-section, and longi- 5.0 60% 45% 75 3
tudinal cross-section.
5.25
52% 65% 120 6
(JA milk line)
R5 occurs approximately 31 to 33 days after R1 (Figure 6). 5.5
40% 90% 175 10
Plants defined as R5 have kernels that are "dented" at t he (Y2 milk line)
kernel top due to declining moisture content and increas- 5.75
37% 97% 205 14
ing starch content. Kerne l moisture is approximat ely 60% (-%milk
at the beginn ing of R5. 65 The "mi lk line" is the zone of 6.0
separation between the softer doughy white portion (Physiological 35% 100%
nearest the cob and t he starchy sol id portion at the top.
Ears at R5 have husk leaves t hat are fading to a pa le green TOTAL
575 33
and browning on the edges. (AVERAGE)

Staging kernels within R5 is possible simp ly by identifica-


tion of the milk line on the non-embryo side of the kerne l
or by slicing the kernel longitudina lly and looking inter-
nally. The sta rchy solid interior portion moves from t he top
of the kernel toward the cob as the kernel matures. Most
often, kernels w ith in R5 are specifical ly designated by t he
progression of t he mi lk line: 1,4, Y2 , or%. Observi ng t he
milk line functions as a good, fie ld-based too l to estimate
kernel deve lopment 67 although it is not di rectly correlated
to dry matter accumu lation 68 Progression of t he m il k
line and t ime req uired betwee n each quarter varies due
to temperature, 69 available moisture, and hybrid matu-
rity68 w ith the expected trend shown in Table 3. The time
needed to reach R6 from the % mi lk line is significantly
greater than between the other quarter mi lk line positions
(Figure 6 and Table 3).

Kerne l dry matter accumu lation is approximately 45% of


tota l dry weight at the beginning of R5, leaving more t han
half to be accumulated during th is last stage. Once kernels
have reached 5.5 (Y2 m ilk line), approximately 90% of total
dry matter exists 67·68 · 69 Environmental stress occurring
during R5 results in a reduction of carbohydrates provided
from the plant, resu lting in reduced kernel weight.

Figure 74. Primary ear at RS shown with and without husk leaves and
silks. The base of the shank is the point of attachment to the stalk.

38 Reproductive Stages
Figure 75. Three uppermost ear shoots of RS Figure 76. Three uppermost ears with silks; primary Figure 77. Three uppermost ears without silks;
plant; primary ear on right. ear on right. primary ear on right.
•.

Figures 75, 76, and 77. Three uppermost ear shoots and ears from RS plant. The change in silk color to brown marks the transition area from inside to
outsi de the husk leaves for each silk. The secondary ear and silks are beginning to rot and turn brown.

Reproductive Stages 39
R6: REPRODUCTIVE STAGE 6
(PHYSIOLOGICAL MATURITY)

I I '
" v t
Figure 78. Kernel s from R6 plant. Kern els are arranged L to R:
whole on embryo side, whole on non-embryo side, pl anar
cross-section , and longitudinal cross-section .

R6 occurs approximately 64 to 66 days after Rl,


with half of this time spent in stageRS (Figure 6
and Table 3). Plants defined as R6 have kernels
at physiolog ica l maturity (PM). Kernels at R6 no
longer have a milk line as that has progressed
completely down to the cob (Figure 78), and have
maximum dry matter (Figure84) . Kerne l moisture
at PM is approximately 35% with a range of at
least +/-2% due to hybrid genetics and environ-
ment.ll. 72 Leaves and stalk tissue are green-brown
at R6 with green tissue becom ing less (Figure 79)
as moisture content of the plant decreases.

Following PM, an abscission layer (comprised of


accumulated carbon) forms at the kernel base
eliminating further dry maner accumulation; this
is referred to as the "black layer:' Physiological
maturity and black layer (BL) are terms often used
interchangeably although they are not the same.
It is not possible to visual ly identify the exact
point in time when a hybrid has reached PM,
wh ich is one primary reason the black layer is
used and recommended often in determ ining
hybrid maturity. 73 Formation of the BL is not
instantaneous and can be visual ly tracked as
it progresses from light grey to dark brown to
black_74 · 75 The grain moisture associated with BL
is much more d ifficult to predict than PM and
wi ll vary widely although a general range is 28%
+/-4%. 75· 76 Environmental factors, specifical ly
stresses li ke temperature, drought, or disease, can
cause premature formation of the black layerl 6

Figure 79. R6 plant. Lower leaves have dropped and all


leaves are browning . Roots are decomposing and the root
mass deteriorating (compare to Figure 57).

40 Reproductive Stages
Final kernel weight varies significantly due to environment and hybrid. Average
kernel weights are approximately 350 mg per kernel (at 15.5% moisture) but
can range from 200 to 430 mg per kernel. 66 Therefore, assuming 56 pounds of
grain are equal to a bushel, this equates to 73,000 kerne ls per bushel (average)
and a range of 59,000 to 127,000 kernels per bushel 66 Kernel we ights toward
the upper portion of the range reflect an environment very conducive to grain
fi ll. An ear w ill typical ly have 450 to 550 kernels total based on recommended
practices and a favorable environment 62

Grain moisture continues to decrease after R6 at a near linear rate w ith reduc-
tions of approximately 0.5 to 0.75% per day until near 20% 65· 77 Environmental
stress occurring after R6 wil l not result in reduced grain yield because kernel
weight is constant. 77 Following physiological maturity, gra in yie ld can only
decrease in events during which the plant or ear is damaged such as stalk
lodging from high winds or feeding by insects or animals.

The amount of grain harvested is the product of the interaction of three factors
(G x M x E): genetic (G) potential of the hybrid, management (M) of the field,
and environmenta l (E) conditions over the course of the growing season.
These three factors influence fina l grain yield by affecting the following yie ld
components to varying degrees: (1) number of plants per acre, (2) ears per
plant, (3) kernel rows per ear, (4) kernels per row, and (5) we ight of individual Figure 80. Primary ear at R6 shown with and
kerne ls. In general, th ese components are determined sequentia lly progressing without hu sk leaves and silks. The base of the
shank is the point of attachment to the stalk.
during th e growing season in the order they are listed.

·.

Figure 81. Two uppermost ear shoots of R6 plant; Figure 82. Two uppermost ears with silks; primary Figure 83. Two uppermost ears without silks;
primary ear on right. ear on right. primary ear on right.

Figures 81, 82, and 83. Two uppermost ear shoots and ears from R6 plant. The secondary ear and silks are rotting, resulting in the brown coloration.

Reproductive Stages 41
The rate of increase and the amount of dry
Dry Matter Accumulation matter (DM) accumulated during the growing
season differs based on the plant component
12600 14.1
measured (Figure 84). Yet when all components
are summed together, it genera lly follows an
50 10500 11.8 S-shaped response curve (Figure 85). More than
20,000 lb/acre (22,400 kg/ha) of above-ground
l:
Cll DM is produced in fields if given ample sun light,
~ 40
~
8400 !..

.
!!!u
9.4 !..
..
.a, wate r, and nutrients wit hout compet ition from
c
~
1-
30
!..
6300 .s::.
Cll
7. 1
~
i
weeds, insects, or diseases.
'iii 'iii
~ ~
~ ~
The sta lk an d leaf sheath s contain the greatest
4200 c 4 .7 c amount of DM among the vegetative co mpo-
nents, w ith maximum weight reached at R2.
10 2100 2.3 Th e amount of stalk and leaf sheath DM beg ins
to decline afterwards, presumably due to nutrient
reallocation from the sta lk to the developing
0
0 500 1000 1500 2000 2500 ear. The leaf blades are at maximum dry matter
GOOF
at R2 and th is is unchanging until th e plant nears
E 3 6 9 14 18
Vegetative (V) Stage R6 and the leaves senesce. The rate of increase
2 3 4 5 6 in total DM is generally consistent prior to R2 as
Reproductive (R) Stage
we ll as after R3 (Figure 85). Th e lack of substantial
Figure 84. Dry matter (OM) accumulation of individual above-ground components. increase in DM during R2 is expected due to
the comp letion of vegetative development and
21000 23.5
tran sition to kernel development occuring.

The harvest index (HI) of a crop is determined


80 16800 18.8
by dividing the amount of gra in dry matter by
l:
Cll
'iii
~
.e- .
!..
the total plant dry matter (a ll above-ground dry
~ 60 12600 u 14 .1 .a, matter including grain) at phys iologica l maturity.
c
~
0
!l: ~
l:
Corn hybrids grown currently have an approx-
...
1- Cll Cll imate HI of 0.5 with some evidence that thi s
~
0 'iii
- 40 6400 ~ 9.4 may be increasing due to breeding efforts w hen
~ ~ ~
c
c.. c compared with older hybrids9 · 18• 19 Regardless
..!:!
0..
of the exact HI va lue, it is clear that as producers
20 4200 4 .7
strive for increased grain yie lds from an acre
of land, the amount of vegetative dry matter
increases by a similar amount. Sig nificant dry
0 0
500 1000 1500 2000 2500 matter accumulation w ithin the grain does not
GOD,
occur prior to R2, although a rapid and near
E 3 6 9 14 18
Vegetative (V) Stage linear increase occurs from R2 to approximately
2 3 4 5 6 R5. 75 (% milk line) (Figure84).
Reproductive (R) Stage

Figure 85. Cumulative dry matter (OM ) of above-ground components .

Figures 84 and 85. Individual and cumulative dry matter (OM ) accumulation, * Quantity of dry matter, lb/acre and Mg/ ha, can vary from values
on a percentage and pound-per-acre basis from VEto R6. Grain yield averaged shown due primarily to management and environment.
225 bu/acre (14.1 Mg/ ha). They-axis scales(%, lb/acre, and Mg/ ha) change
between the figures. Approximate vegetative and reproductive stages are
shown on the x-axis for reference .

42 Dry Matter Accumulation


Nutrient Accumulation
Three macronutrients account for the majority of
fertilizer applied to meet crop demand: nitrogen
(N), phosphorus (P), and potassium (K). Although
other macro- and micronutrients are needed,
they are not regularly applied by producers
due to the relatively sma ll amount needed and
adequate available natural levels. The rate of
increase and the amount of each nutrient (N, P.
and K) accumulated during the growing season
differs based on the plant component measured.
Yet when all components are summed and total
N (Figure87), P (Figure89), and K (Figure 91) are
viewed, they follow a looseS-shaped response
curve. Each nutrient is reported here in elemental
form and wi ll need to be converted if wanting
to determine the equivalent contained within
fertilizer formulations, such as P20 5 and K20.
Variation is expected to occur with N, P, and K
accumulation among hybrids, management
systems, and years. The following figures serve
as an average for most environments.

Sign ificant nutrient accumulation within the


grain does not occur prior to R2, although a
rapid and near linear increase begins at R2 to
approximately RS. During RS, N and P continue
to be accumulated at a similar rate as before
but K accumu lation is lessened.

At physiological maturity, the plant contains


nearly 200 lb N/acre (224 kg/ha) (Figure 87). Total
plant phosphorus and potassium is less at 36
and 125 lb/acre (40 and 139 kg/ha), respective ly
(Figures 89and 91).

Nutrient Accumulation 43
NITROGEN (N)
Except for gra in, leaf blades have the highest fraction of N (75 lb N/acre (84 kg N/ha)) with maximum accumu lation at
R2 (Figure 86). Components decrease in total nitrogen at R1 or R2 due to remobilization of the nitrogen to the developing
grain as we ll as senescence of the lower leaves. When mature (R6), 67% of total plant nitrogen, or nearly 140 lb N/acre
(157 kg N/ha), is contained in the grain (Figure 86).

157
Figure 86. Nitrogen (N) accumulation
of individual above-ground components.
60 120 134

c 50 100 !- 112 .
li
....f!
Cll
C) .s::.
~ ~
z
z
~
C)
40 80 @. 90 ~
!! Cll Cll
...:
~ ...:
!! !!
0 c. c.
~ 30 60 ::I 67 ::I
c
~
c Cll
....c Cll
C)
C)

Cll
.... .~ ~ - Stalk and Leaf Sheaths
~ 20 40 z 45 z
0.. - Leaf Blades

- Tassel
10 20 22
- Shank, Husk Leaves, and Cob

0 0 - Grain
0 500 1000 1500 2000 2500
GDDF
E 3 6 9 14 18
Vegetative (V) Stage
2 3 4 5 6
Reproductive (R) Stage

200 224
Figure 87. Cumulative nitrogen (N) accumulation
of above-ground components.

80 160 179

c !-
!-
....f!
Cll

z"'
C)
.s::.
~ ~
z 60 120 z
@.
134
~
C)

~ Cll Cll
...:
0 ...:
...
1-
0 a
::I
!!
c.
::I Figures 86 and 87. Individual and cumulative
~ 40 80 c 90 c
Cll nitrogen (N) accumulation on a percentage
....c Cll
C)
0
C)
0
Cll !::: :2 and pound-per-acre basis from VEto R6 . Grain
l:!
Cll
0..
z z yield averaged 225 bu/acre (14.1 Mg/ ha) . The
y-axis sca les (%, lb/acre, and kg /ha) change
20 40 45
between the figures. Approximate vegetative
and reproductive stages are shown on the
x-axis for reference .
0 0
0 500 1000 1500 2000 2500
GOOF
E 3 6 9 14 18 *Quantity of nitrogen, lb/acre and kg / ha, can vary
Vegetative (V) Stage
from values shown due primarily to management
2 3 4 5 6 and environment.
Reproductive (R) Stage

44 Nutrient Accumulation
PHOSPHORUS (P)
All components, except fo r t he tasse l and gra in, contain nea r equa l fractions of phosphoru s with maxi mum accu m ulation
at approximately R2 (Sib P/acre (9 kg P/ha)) (Figure 88). Components decrease in total phosphorus at R2 due to remobil i-
zation of the phosphorus to the developing gra in as wel l as senescence of the lower leaves. When mature (R6), approxi-
mately 80% of tota l plant phosphorus, o r 30 lb P/acre (34 kg P/ha), is contai ned in the gra in (Figure 88) .

40
Figure 88. Phosphorus (P) accumu lation
of ind ividual above-ground components .

75 27 !-... 30 •
VI
:::J ~ <V
u .s=
0
.s=
IU
0::
c. 0:: Cl
VI
0
.s=
~ ~
II>
a. II>
-" -"
IU

~ 50 18 -5. 20 Q.
::I
...
1-
0
::I
VI
:::J
VI
:::J

~ 0 0c.

-
L
c:
II>
::II> 25 9
.s=
c.
VI
0
.s=
a. 10 a.
.s=
VI
0
.s=
-

-
Stalk and Leaf Sheaths

Leaf Blades
a.
- Tassel

- Shank, Husk Leaves, and Cob

0 0 0 - Grain
0 500 1000 1500 2000 2500
GOOF
E 3 6 9 14 18
Vegetative (V) Stage
2 3 4 5 6
Reproductive (R) Stage

100 36 40
Figure 89. Cumulat ive phosp horus (P)
accumulation of above-ground components .

VI 75 27 !-... 30 •
:::J ~ <V
0 u .s=
.s= ~ 0::
c.
VI
a. Cl
0
.s= ~ ~
a. II> II>

...-
-" -"
IU
iii IU

1-
0
0 50 18 Q.
::I
VI
20 Q.
::I
VI
:::J
.
Figures 88 and 89. Individual an d cumulative
:::J
0c.
~ 0
.s=
c. .s=
phosphorus (P) accum ul ation on a percentage
cII>
VI
0
VI
0
and pound -per-acre basis from VEto R6. Grain
u .s= .s=
:D 25 9 a. 10 a. yield averaged 225 bu/acre (14.1 Mg/ha).
a. Approximate vegetative and re produ ctive
stages are shown on the x-axis fo r reference .

0 0 0
0 500 1000 1500 2000 2500
GOOF
E 3 6 9 14 18 * Quantity of phosphorus, lb/acre and kg/ha, ca n
Vegetative (V) Stage
varyfrom values shown due primarily to management
2 3 4 5 6
Reproductive (R) Stage
and environment.

Nutrient Accumulation 45
POTASSIUM (K)
Potassium uptake occurs rapidly during the early vegetative stages with the majority of total plant K within the leaf
blades and the stalk and leaf sheaths. Maximum accumulation of K with in these two components occurs at Rl or R2
(Figure 90) and total plant K is nearly 90% by R2 (Figure 91). The slight fluctuation of K in the sta lk and leaf sheaths after
R2 is presumably due to remobilization of K withi n the plant. Al l components, except for gra in, genera lly decrease in
tota l K after Rl or R2; this is related to remobilization within the plant and/o r leaching from lower leaves that have
senesced. Potassium is highly mobi le w ith in the plant, creating greater flu ctuation w ithi n individual co mponents than
observed with Nor P; thi s is similar to previous research. 6 When mature (R6), about 30% of total plant potassium, or
approximately 40 lb K/acre (45 kg K/ ha), is conta ined in th e grain (Figure 90).

76
Figure 90. Potassium (K) accum ulatio n
of ind ividual above-groun d compon ents .

E 37.5 51
:.... 57
.
.=
·;;;
.'!
.
I!!
u
;z
li
..c:
;z
0 Cll
D. @. ~

~ 01> 01>

...
1- 25 34 ""'i 38 .'!""'
ca.
0 :;) :;)

:.!! E E

•.'!.= .=
0

'iii
cQl - Stalk and Leaf Sheaths
~
.'!
0 0
Ql
D. D.
D. 12.5 17 19 - Leaf Blades

- Tassel

- Shank, Husk Leaves, and Cob

0 0 0 - Grain
0 500 1000 1500 2000 2500
GOOF
E 3 6 9 14 18
Vegetative (V) Stage
2 3 4 5 6
Reproductive (R) Stage

100 136 152


Figure 91 . Cumulative potassi um (K) accumulation
of above-grou nd components.

E 75 102. 114 .
.=
'iii !u
..
;z
li
..c:
;z
~ Cll
D. @. ~
! 01> J
::. 50 68 ""'i 76 .'!
ca. Figures 90 and 91. Individual and cumulative
0 :;) :;)

..~
c
Ql
E

..= ~..=
·;;
E
'iii
potassi um (K) accumulation on a percentage
an d pou nd-per-a cre bas is from VE to R6. Grai n
~ .'! yield averaged 225 bu/a cre (14. 1 Mg/ ha) . Th e
Ql 0
D.
25 34 0..
D. 38 y-ax is scal es (o/o, lb/acre, and kg/ha) change
between the fi gures. Approximate vegetative
and re pro ductive sta ges are shown on the x-axis
fo r refere nce.
0 0
0 500 1000 1500 2000 2500
GOOF
E 3 6 9 14 18 * Quantity of potassium, lb/acre and kg/ha, can
Vegetative (V) Stage vary from values shown dueprimarily to management
2 3 4 5 6 and environment.
Reproductive (R) Stage

46 Nutrient Accumulation
14
0tegui, M.E. and R. Bonhomme. 1998. Grain yield components in maize:
ENDNOTES I. Ear growth and kernel set. Field Crops Res. 56:247-256.
The scientific literature cited here represents on ly a small
15
fraction of t he agronomic research publ ished re lative to Borras, L., M.E. Westgate, J.P. Astini, and L. Echarte. 2007. Coupling time to
corn growth and development. This is not intended to be silking with plant growth rate in maize. Field Crops Res. 102:73-85.
all inclusive of existing research literature.
16
USDA-National Agricultural Statistics Service. 2010. Corn: Acreage by Year, US.
1
Ritchie, S.W., J.J. Hanway, and G.O. Benson. 1986. How acorn plant develops. Available at www.nass.usda.gov/Charts_and_Maps/Field_Crops/cornac.asp.
Spec. Rep. 48.1owa State Univ. Coop. Ext. Serv., Ames, Iowa. USDA-NASS, Washington, D.C.

17
2
Hanway, J.J. 1966. How acorn plant develops. Spec. Rep. 48. 1owa State Univ. FAOSTAT, Statistics Division, Food and Agriculture Organization of the
Archives. United Nations. 2008. Top Production-World (Total)-2008. Available at
faostat.fao.org/site/339/default.aspx.
3
Hanway, J.J. 1963. Growth stages of corn (lea mays, L.). Agron. J. 55:487-492. 18
Tollenaar, M., and E.A. Lee. 2002. Yield potential, yield stability and stress
4
Frank, A.B., V.B. Cardwell, A.J. Ciha, and W.W. Wilhelm. 1997. Growth staging tolerance in maize. Field Crops Res. 75:161-169.
in research and crop management. Crop Sci. 37:1039-1040. 19
Duvick, D.N., J.S.C. Smith, and M. Cooper. 2004. Long-term selection in a
5
Hanway, J.J. 1962a. Corn growth and composition in relation to soil fertility: commercial hybrid maize breeding program. p. 109-151./n J. Janick (ed.)
I. Growth of different plant parts and relation between leaf weight and grain Plant breeding reviews. Vol. 24, Pt. 2. John Wiley &Sons, N.Y.
yield. Agron. J. 54:145-148. 20
Tollenaar, M. and J. Wu. 1999. Yield improvement in temperate maize is
6
Hanway, J.J. 1962b. Corn growth and composition in relation to soil fertility: attributable to greater stress tolerance. Crop Sci. 39:1597-1604.
II. Uptake of N, P, and Kand their distribution in different plant parts during the 21
growing season. Agron J. 54:217-222. Duvick, D.N. and K.G. Cassman. 1999. Post-green revolution trends in yield
potential of temperate maize in the north-central United States. Crop Sci.
7
Syngenta Seeds, Inc., Minneapolis, Minn. 55440. Golden Harvest®is aregistered 39:1622-1630.
trademark of Golden Harvest Seeds, Inc.
22
Tollenaar, M., and E.A. Lee. 2006. Dissection of physiological processes
8
The Corn Belt is adescriptor for aregion, or collection of states in the U~ited underlying grain yield in maize by examining genetic improvement and
heterosis. Maydica 51 :399-408.
States, that produces asignificant amount of corn. States with significant corn
acreage are documented by the USDA-NASS, see: USDA-National Agricultural 23
Statistics Service. 2007. Ag Atlas Maps: Corn Grain, Harvested Acres: 2007. Avail- Aitered root architecture is substantiated with crop models only and not
able at www.agcensus.usda.gov/Publications/2007 /Online_Highlights/ Ag_At- in-field quantitative data; see the following paper for analysis: Hammer,
las_Maps/Crops_and_Piants/index.asp. USDA-NASS. Washington, D.C. G.L., Z. Dong, G. Mclean, A. Doherty, C. Messina, J. Schussler, C. Zinselmeier,
S. Paszkiewicz, and M. Cooper. 2009. Can changes in canopy and/or root
9
Boyer, M.J. 2011 . Dry matter and nutrient uptake in maize hybrids from the system architecture explain historical maize yield trends in the U.S. Corn Belt?
Crop Sci. 49:299-312.
1960'sto 2000'sin central Iowa. M.S. thesis. Iowa State University, Ames, Iowa.
24
10
Pioneer Hi-Bred International, Inc. Johnston, Iowa. Available at Fehr, W.R., C.E. Caviness, D.T. Burmood, and J.S. Pennington. 1971. Stage
www.pioneer.com. of development descriptions for soybeans, Glycine max (L.) Merrill. Crop Sci.
11 :929-931 .
11
USDA-National Agricultural Statistics Service. 2007 and 2008. Annual 25
Crop Summary. Available at www.nass.usda.gov/Statistics_by_State/lowa/ Fehr, W.R. and CE. Caviness. 1977. Stages of soybean development. Iowa
Publications/Crop_Report/. USDA-NASS. Washington, D.C. Agric. Exp. Stn. Spec. Rep. 80. Iowa Agric. Home Econ. Exp. Stn., Iowa State Univ.,
Ames, Iowa.
12
Sawyer, J., E. Nafziger, G. Randall, L. Bundy, G. Rehm, and B. Joern. 2006. 26
Concepts and rationale for regional nitrogen rate guidelines for corn. PM 2015. Text, content, and design were modified from the original figure to primarily
Iowa State Univ. Coop. Ext. Serv., Ames, Iowa. aid in comprehension. Figure as shown here has been approved by G.S. McMas-
ter for publication. The original figure (Figure 5 on page 1283) is published in
13
Systat Software, Inc. 2006. Sigma Plot for Windows. Version 10.0. Systat McMaster, G.S., W.W. Wilhelm, and A.B. Frank. 2005. Developmental sequences
Software, Inc. Point Richmond, Calif. for simulating crop phenology for water-limiting conditions. Aust. J. Agric. Res.
56:1277-1288.

Endnotes 47
27 41
Abendroth, L.J. and R.W. Elmore. Unpublished leaf number data (n=875 Abendroth, L.J., S.K. Marlay, and R.W. Elmore. Unpublished emergence data
measurements) collected from Iowa State University multi-location planting (n=675 measurements) collected from three Iowa State University research
date research conducted from 2007 to 2009. trials conducted near Ames, Iowa in 2009 and 2010.

28 42
USDA-Federal Crop Insurance Corporation. 2010. Corn loss adjustment Abendroth, L.J., A.J.W. Myers, M.J. Boyer, S.K. Marlay, and R.W. Elmore. Graphic
standards handbook. Available at www.rma.usda .gov/handbooks/25000/ derived from unpublished vegetative (n=600 measurements) and reproductive
201 0/10_25080.pdf. USDA-FCIC. Washington, D.C. (n=200 measurements) developmental data staged to 0.25 accuracy. Data col-
lected from three Iowa State University research trials conducted from 2007 to
29
Staging method as used and described in Tollenaar, M., T.B. Daynard, and R.B. 2010 near Ames, Iowa with hybrids ranging in relative maturity from 108 to 112.
Hunter. 1979. Effect of temperature on rate of leaf appearance and flowering
date in maize. Crop Sci. 19:363-366. 43
Number of days based on 15 April planting and 1May emergence (VE) dates.
Day projections were derived using GDD requirements shown in Figures 5
30
BBCH is an abbreviation for the BASF-Bayer-Ciba-Geigy-Hoechst method and (vegetative) or 6 (reproductive) paired with 20-year Ames, Iowa weather data
the methodology is published in Lancashire, P.D., H. Bleiholder, T. Van Den Boom, (1990-2009) from the Iowa Environmental Mesonet (www.mesonet.agron.
P. LangeiOddeke, R. Stauss, E. Weber, and A. Witzenberger. 1991 . Auniform deci- iastate.edu) and +/-1 day added to each projected date.
mal code for growth stages of crops and weeds. Ann. Appl. Bioi. 119:561-601.
44
McMaster, G.S., W.W. Wilhelm, and A.B. Frank. 2005. Developmental sequences
31
Harrell, D.M., W.W. Wilhelm, and G.S. McMaster. 1998. SCALES 2: Computer for simulating crop phenology for water-limiting conditions. Aust. J. Agric. Res.
program to convert among developmental stage scales for corn and small 56:1277-1288.
grains. Agron. J. 90:235-238.
45
Nielsen, R.L., P.R.Thomison, G.A. Brown, A.L. Halter, J. Wells, and K.L.
32
Abendroth, L.J. and R.W. Elmore. Unpublished vegetative developmental Wuethrich. 2002. Delayed planting effects on flowering and grain maturation
data (n=400 measurements) staged to 0.25 accuracy in two hybrids (110 and of dent corn. Agron. J. 94:549-558.
112 relative maturity) using multiple staging and height methodologies. Iowa
46
State University research conducted near Ames, Iowa in 2009. Markelz, N.H., D. E. Costich, and T.P. Brutnell. 2003. Photomorphogenic
responses in maize seedling development. Plant Phys. 133:1578-1591 .
33
Muldoon, J.F., T.B. Daynard, B. Van Duinen, and M. Tollenaar. 1984.
47
Comparisons among rates of appearance of leaf tips, collars, and leaf area Singh, V., E.J. van Oosterom, D.R. Jordan, C. D. Messina, M. Cooper, and G.L.
in maize (lea mays l.). Maydica 29: 109-120. Hammer. 2010. Morphological and architectural development of root systems
in sorghum and maize. Plant Soil. 333:287-299.
34
Abendroth, L.J, A.J.W. Myers, S.K. Marlay, and R.W. Elmore. Unpublished stalk
48
internode length data (n= 120 measurements) collected from three planting Hochholdinger, F., K. Woll, M. Sauer, and D. Dembinsky. 2004. Genetic
date research projects conducted at Iowa State University near Ames, Iowa in dissection of root formation in maize (lea mays) reveals root-type specific
2009. Data reported only from plantings in early May. developmental programmes. Ann. Bot. 93:359-368.

35 49
Russell, W.A. 1985. Evaluations for plant, ear, and grain traits of maize cultivars Hoppe, D.C., M.E. McCully, and C.L. Wenzel. 1986. The nodal roots of Zea:
representing seven eras of breeding. Maydica 30:85-96. their development in relation to structural features of the stem. Can. J. Bot.
64: 2524-2537.
36
Tollenaar, M. 1991 . Physiological basisof geneticimprovement of maize
50
hybrids in Ontario from 1959-1988. Crop Sci. 31 :119-124. liedgens, M., A. Soldati, P. Stamp, and W. Richner. 2000. Root development
of maize (lea mays L.) as observed with minirhizotrons in lysimeters.
37
By staging plants as R1 under this scenario, it places more priority on the Crop Sci. 40:1665-1672.
presence of silks rather than the exsertion of all tassel branches. This change in
51
staging reflects the importance of silking as an indicator of overall agronomic lied gens, M. and W. Richner. 2001 . Minirhizotron observations of the spatial
yield, more than tassel branch emergence. distribution ofthe maize root system. Agron. J. 93:1097-1104.

38
Stewart, D.W., L.M. Dwyer, and L.L. Carrigan. 1998. Phenological temperature 52
.
Dardanell i, J.L., O.A. Bachmeier, R. Sereno, and R. Gil. 1997. Rooting depth
response of maize. Agron.J. 90:73-79. and soil water extraction patterns of different crops in a silty loam Haplustoll.
Field Crops Res. 54:29-38.
39
Dwyer, L.M., D.W. Stewart, L. Carrigan, B.L. Ma, P. Neave, and D. Balchin. 1999.
53
Ageneral thermal index for maize. Agron. J. 91 :940-946. Summary of multiple research papers that are listed in Table 1of Borg, H.
and D.W. Grimes. 1986. Depth development of roots with time: An empirical
40
Cross, H.Z. and M.S. Zuber. 1972. Prediction of flowering dates in maize based description. Trans. ASA£29(1):194-197.
on different methods of estimating thermal units. Agron. J. 64:351 -355.

48 Endnotes
54 68
McDonald, M.B., J. Sullivan, and M.J. Lauer. 1994. The pathway of water Ma, B.L. and L.M. Dwyer. 2001 . Maize kernel moisture, carbon and nitrogen
uptake in maize seeds. Seed Sci. Techno/. 22:79-90. concentrations from silking to physiological maturity. Can. J. Plant Sci.
81:225-232.
55
Miedema, P.1982. The effects of low temperature on lea mays. Adv. Agron.
69
35: 93-128. Muchow, R.C. 1990. Effect of high temperature on grain-growth in field-grown
maize. Field Crops Res. 23:145-158.
56
Kucharik, C.J. 2006. Amultidecadal trend of earlier corn planting in the
70
Central USA. Agron. J. 98:1544-1550. Th is table is a compilation of several published research papers and is the
expected trend for RS with some variation. Research used in making this table
57
Kiesselbach, T.A. 1999.The structure and reproduction of corn. 50th includes Figure 84 and endnotes 67, 68, 69, and 71.
Anniversary Edition. Cold Spring Harbor Lab. Press, Cold Spring Harbor, N.Y.
71
Borras, L., C. Zinselmeier, M.L. Senior, M.E. Westgate, and M.G. Muszynski.
58 2009. Characterization of grain-filling patterns in diverse maize germplasm.
Stevens, S.J., E.J. Stevens, K.W. Lee, A.D. Flowerday, and C.O. Gardner. 1986.
Organogenesis of the staminate and pistillate inflorescences of pop and dent Crop Sci. 49:999-1009.
corns: Relationship to leaf stages. Crop Sci. 26: 712-718.
72
Sala, R.G., F. H. Andrade, and M.E. Westgate. 2007. Maize kernel moisture at
59
Abendroth, L.J. and R.W. Elmore. Unpublished ear node data (n= 175 measure- physiological maturity as affected by the source-sink relationship during grain
ments) collected from Iowa State University planting date research conducted filling. Crop Sci. 47:711-716.
near Ames, Iowa in 2009.
73
Daynard, T.B. 1972. Relationships among black layer formation, grain moisture
60
Carcova, J., B. Andrieu, and M.E. Otegui. 2003. Silk elongation in maize: percentage, and heat unit accumulation in corn. Agron. J. 64:716-719.
Relationship with flower development and pollination. Crop Sci. 43:914-920.
74
Rench, W.E. and R.H. Shaw. 1971. Black layer development in corn. Agron. J.
61 63:303-305.
Uribelarrea, M., J. Carcova, M.E. Otegui, and M.E. Westgate. 2002. Pollen pro-
duction, pollination dynamics, and kernel set in maize. Crop Sci. 42:1910-1918.
75
Hunter, J.L., D.M. TeKrony, D.F. Miles, and D.B. Egli. 1991 . Corn seed maturity
62 indicators and their relationship to uptake of Carbon-14 assimilate. Crop Sci.
Abendroth, L.J. and R.W. Elmore. Unpublished total kernel number/ear data
(n=80 measurements) collected from Iowa State University research conducted 31:1309-1313.
near Ames, Iowa in 2008. -.
76
Afuakwa, J.J., R.K. Crookston, and R.J. Jones. 1984. Effect oftemperature and
63 sucrose availability on kernel black layer development in maize. Crop Sci.
Westgate, M.E., M.E. Otegui, and F.H. Andrade. 2004. Physiology of the corn
plant. in C.W. Smith, J. Betran, and E.C.A. Runge. Corn: Origin, History, Technol- 24:285-288.
ogy, and Production. John Wiley &Sons, Inc. Hoboken, N.J.
77
Eimore, R.W. and F.W. Roeth. 1999. Corn kernel weight and grain yield stability
64 during post-maturity drydown.J. Prod. Agric. 12:300-305.
Bassetti, P.and M.E. Westgate. 1994. Floral asynchrony and kernel set in maize
quantified by image analysis. Agron. J. 86:699-703.

65
Kernel moisture listed for R2, R3, R4, and RS were derived by aligning grain dry
matter data (as shown in Figure 84) and approximating the respective moisture
from Figure 2A of Borras, L. and B.L. Gambfn. 2010. Trait dissection of maize
kernel weight: Towards integrating hierarchical scales using aplant growth
approach. Field Crops Res. 118:1-12.

66
Kernel weight and the number of kernels/bushel were derived by alterations
to data from Figure 1Bof Borras, L. and B.L. Gambfn. 2010. Trait dissection
of maize kernel weight: Towards integrating hierarchical scales using a plant
growth approach. Field Crops Res. 118:1-12. Kernels were oven dried in the
original paper and these were approximated to be at 3.0%moisture content
to allow for conversion to 15.5% moisture content.

67
Afuakwa, J.J. and R.K. Crookston. 1984. Using the kernel milk line to visually
monitor grain maturity in maize. Crop Sci. 24:687-691 .

Endnotes 49

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