Documenti di Didattica
Documenti di Professioni
Documenti di Cultura
191
.M2
C678
2011
Photography by
Iowa State University
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 :
:Internode Elongation :
-
·~
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
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 ....... ...... .
·~
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
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 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 .
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.
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).
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
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.
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
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.
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).
20 Vegetative Stages
Figure 26. Dissected V12 plant. Figure 27. Tassel from a V12 plant.
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 ).
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
.. ....
f
I rSh•nk
a- 0 Kernels
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
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
••
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
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
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.
Figure 58. Glumes partially surrounding each kernel are visible when
looking at an ear cross-section .
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 .
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 .
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)
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 .
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.
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 .
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
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
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
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