Responses of Maize to Plant Population Density. I.

Canopy Development, Light Relationships, and Vegetative (Growth

F. Tetio-Kagho and F. P. Gardner*
ABSTRACT
Understanding of maize (Zea mays L.) canopy structure and light transmission over a wide range of plant population densities (PPD) is necessary in the formulation of maize intercrop associations. Experiments using a systematic (fan) design were conducted on Lake fine sand (hyperthermic, coated Typic Quartzipsamments) at Gainesville, FL (2938N), 1985 and 1986 to assess the interrein lationships among maize canopy structure, light interception, and vegetative growth over 15 PPD ( . to 15.4 plants m-2). Fractional 08 light transmission (by canopy levels at different times up to noon), leaf area index (LAI), total dry matter (TDM) yield, plant height, and tiller number were recorded. Light interception was similar at 5 and 3 h before and at solar noon for each PPD measured at ground, ear, and below-tassel levels. Leaf area and light interception were highly concentrated at ear level, but level of light interception shifted upward with increasing PPD. Maxima LA1 were 17 2 6 and 4 0 ., . , . at tasseling for the 17 2 6 and 6.3 plants m-: respectively. Light ., . , interception by tassels was approximately 2 30, and 40% for 1 9 , ., 3 5 and 6 3 plants m-*, respectively, while that for whole canopy ., . was 75, 90, and 97%, respectively. As early as 35 d after planting (DAP) canopy interception was 40, 60, and 75% for 1 9 3 5 and . , ., 6.3 plants m-z, respectively. Leaf area index, TDM, crop growth rate, and plant height were significantly influenced by PPD. Tiller number decreased linearly with increasing PPD to no tillers at 3 5 . plants m-*. We conclude that increasing PPD of maize increases LAP and vegetative DM yield but alters light distribution within the canopy by shifting it from lower to upper canopy strata and increasing fraction intercepted by tassels.

canopy is the large attenuation of light in the top 50 cm of the canopy at high PPI) (>50 000 plants hair1), a critical factor in canopy illumination (Loomis et al, 1968). Duncan (1967) demonstrated that at high EPD light interception by tassels ranged from approximately 4 to 20% incident irradiance over the range of 17 500 to 125 000 plants ha-, which reduced light availability to the more photosynthetic leaf surfaces. Generally, studies relating growth functions to IPD in maize have employed conventional rows and relatively narrow PPD ranges. Also PPD was confounded with plant spatial arrangement, since row widths were constant, and PPD was varied only by adjusting distance between plants in the row, which altered spatial arrangement. Little information is available on effects of PPD on canopy structure where plant spacing was equidistant or gridded despite the fact that such arrangement is commonly used in the tropics for maize intercropping. This study was designed to assess the relationships of canopy Structure, light interception, and vegetative growth of maize at low to ultra-high PPD in equidistant spacing. MATERIALS AND METHODS
Field experiments were conducted at Gainesville, FL, in 1985 a n d 1986 on a Lake fine sand. The maize hybrid Pioneer Brand 3192 was planted in a systematic design (fan) (Nelder, 1962) at 15 PPD ranging from 0.80 to 15.4 plants m-* (23.5% increment between PPD). The fan (Fig. 1) contained 29 radii a n d 20 arcs including borders. The dimensions were calculated from Bleasdales (1967) equations for design,

in crop plants is directly related to the utilization of solar radiation (Donald, 1963; Williams et al., 1968; Daughtry et al., 1983), which is influenced by canopy structure. Williams et al. (1968) observed that the effect of canopy architecture on vertical distribution of light within the maize canopy was a major determinant of photosynthetic efficiency and growth. Radiation is transmitted through and between leaves, and its flux density and spectral composition change rapidly with depth (Szeicz, 1974; Gardner et al., 1985). Canopy light interception and photosynthesis were closely related to leaf area index (LAI) up to the critical LAI, that which is required to intercept 95% incident irradiance (Pearce et al., 1965). Williams et al. (1968) found that light interception and crop growth rate (CGR) increased linearly as LA1 increased up to 3, but CGR increased asymptotically as LA1 was increased further to a maximum at 99% light interception. Early et al. (1966) reported that plant height in maize increased, remained static, and decreased as light decreased from 100 to 40%, 30 to 20%, and 20 to 10%of full sunlight, respectively. A particularly striking feature of a maize
RY MATTER PRODUCTION
~~ ~~

r,

3-

4 112 ,

PI

where r,, is the initial distance from the center apex of the fan, A is the smallest area per plant, 0 is the angle in radians

F. Tetio-Kagho, Dep. of Agriculture, Univ. Centre of Dschang, Dschang, Cameroon; and F.P. Gardner, Dep. of Agronomy, Univ. of Florida, Gainesville, FL 32611. Contribution from the Inst. of Food and Agricultural Sciences, Florida Agric. Exp. Stn. Journal no. 8407. Received 8 Oct. 1987. *Corresponding author.
Published in Agron. J. 80:930-935 (1988).

. Fig. 1 Fan design fitted to a rectangular plot as used in 1935 and 1986 experiments.

9 30

TETIO-KAGHO & GARDNER. MAIZE CANOPY STRUCTURE & LIGHT TRANSMISSION

931

between radii, and LY is a constant governing the rate of change. Each other distance (arc) was calculated as follows:

r, = r, a" ; M, to M , = a tan 19, and M, to M,

PI

= a tan 213, [3] where a is the shortest distance from the center of the nearest side (base line) of the enclosing rectangle. The following equations were developed to replace Eq. [3] to enclose fans into rectangular plots (the value of a being too small) to provide additional plants for growth analysis:

puted as the slope of the near linear portion of growth (TDM) versus time. Analyses of variance and regression were used to analyze data using SAS (SAS Institute, 1985) procedures. Differences among treatment means were compared using Duncan's multiple range test (DMRT) or least significant difference (LSD) at the 0.05 probability level.

M , to M,, = M,, = (B M , = L / 2 tan (go

0,
=

+ a) tan On if M,, < L / 2 , [4]

- 0,) - a if M,, > L / 2 , [5] n0. Each fan was replicated three times. The fan design consisted of 29 plants per PPD (arc) with two plants as borders at each arc end. Two arcs at the lowest and three arcs at the highest PPD were added to the 15 PPD for borders. Two maize kernels per hill were hand-planted at each arcradius intersection of the fan on 13 Mar. 1985 and 11 Apr. 1986. A pre-marked vinyl-covered wire was used to delineate the hills at the arc-radius intersection. Hills were thinned to one plant at 21 DAP. Seedbed was prepared by mold-board plowing, tandem disking, and smoothing with a rolling cultivator. A broadcast application of 112-26-100 kg ha-I (N-P-K) was incorporated into the seedbed. Additional fertilizers were applied as side dressing at tasseling as follows: 200 kg N ha-I (",NO,), 45 kg K ha-' (KCl), and 45 kg Mg ha-l (MgSO,). Recommended pesticide applications were made to control weeds, insects, and nematodes, and weeds were also controlled manually. All plots were imgated by a sprinkler system as required (25 mm per application) to alleviate water stress. Three of the PPD (1.9, 3.5, and 6.3 plants m-2) were sampled biweekly, beginning at 15 DAP (V2 stage), for growth analysis in both 1985 and 1986. At early growth stage (V2VS), four plants per treatment in each replicate were harvested for biomass, and two were separated into component parts. After the V9 stage, three plants were harvested, and one was separated into component parts. Measurements included LA1 and leaf, stem, and stalk (leaf plus stem) TDM per plant. A LI-COR' (Model 3100; LI-COR, Inc., Lincoln, NE) leaf-area meter was used to determine leaf area. Leaves and stems were separated and dried at 70C for 2 to 4 d, depending on the stage of maturity of the plant. Light was measured above canopy and at ground level using a LI-COR (Model LI- 188B) integrating recorder and quantum-line sensor (Model 191SB) placed between radii. Light was also measured in the canopy at ground, ear, and below-tassel levels near solar noon (SN) and 5 and 3 h before SN for the 15 PPD at grain-filling period. Percentage light interception (PLI) was determined

RESULTS AND DISCUSSION Light Interception Time o Measurement f Percentage light interception was about the same whether measured at SN or 5 and 3 h before SN for each maize density and canopy level (ground, ear, and below tassel) (Fig. 2, Table 1). This indicates the efficacy of a range of morning times for measurement of canopy light interception. Afternoon measurements were not made since the sun's arc to and from its zenith is the same. However, at PPD of 1.2, 1.5, 1 9 .
Table 1. Regression analysis relating light interception below tassel, at ear level, at ground level, measured at solar noon (SN) and 3 and 5 h before solar noon (SN-3h and SNdh), as a function of maize plant population density (x). Canopy level Measurement time Regression

.. . ....,,,_,.,,.,..,...,.................... E

80 -

-\ " o
z
. . . .. . ...
60-

9 I II
W

'

-SN
I '

SN-3h

PLI

-X 100 -I
la

181

where I, and I are the irradiance above canopy and at a Z) given level in canopy below tassel ( , ,ear level (ZJ, and ground level (I,), respectively. Leaf area distnbution and light interception at various increments of plant height were also recorded. Plant height and tiller number per plant were determined from five central radii for all 15 PPD at physiological maturity on 15 July 1985 and 28 July 1986. Crop growth rate (CGR) was comI Mention of a trademark, vendor or proprietary product is for the benefit of the reader and does not imply endorsement.

IO

12

1 4

1 6

DENSITY (plonts ni2)

Fig. 2 Relationship between light interception and maize plant population density at solar noon (SN) and 3 and 5 h before solar noon ( S N J h and SN-Sh); measurements were made below tassel (T), at ear level (E), and at ground level (G) during grain filling in 1986.

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AGRONOMY JOURNAL, VOL. 80, NOVEMBER-DECEMBER 1988

plants m-2, light interception below the canopy at SN was slightly less than at early hours (5 and 3 h before SN), because the upper, more-erect leaves allowed more light from the midday sun (near normal to the surface) to penetrate the canopy; or conversely, the upper, vertically oriented leaves caused more extinction at lower solar angles.

Efect of Canopy Depth Light interception at SN by the canopy was 83 to 93%at ear level (0.9 m), and in the medium (3.5 plants m-2) and high PPD (6.3 plants m-2) did not decrease interception significantly with greater canopy depth (Fig. 3). Interestingly, at 1.9 plants mu* only 50% of the irradiance was captured above the ear, and interception below the ear increased significantly with canopy depth. These data suggest that in high PPD, necessary for maximum grain yield, light is captured primarily above the ear and by the youngest and more efficient leaves. Approximately 2,30, and 40% of solar radiation was intercepted by tassel at the 1.9, 3.5, and 6.3 plants m-2, respectively, irrespective of solar angle (time measured) (Fig. 2, Table 1). However, measurements at the tassel level were much more variable than those from deeper canopy levels. These high levels of tassel light extinction are consistent with earlier reports (Duncan et al., 1967; Williams et al., 1968). Tassels probably contribute little to photosynthesis. Grogan et al. (1961) reported a positive grain yield response by detasseling, which was also positively related to PPD. Eflect of Plant Population Density Light interception reached a maximum between 77 and 84 DAP for all PPD, which corresponded to the milk and dough (R3 and R4) stages, at a time when LA1 and TDM accumulation rate were maximum. As early as 35 DAP, light interception by the total maize

canopy was 40, 60, and 75% for the 1.9, 3.5, and 6.3 plants m-2, respectively (Fig. 4). These results are important when considering Understory crops in maize intercropping, especially at PPlD greater than 3.5 plants m-* in which only 25% incident light penetrated the canopy as early as 35 DAP. At 63 to 70 DAP (silking to grain filling), 95% incident light was intercepted by both the 3.5 and 6.3 PPD, reflecting canopy closure and maximum interception efficiency for photoaynthesis, but light for understory crop was virtually nil. However, the 1.9 plants m-2 never reached the !)5% interception level; i.e., light penetrated the canopy most of the season. Interception by the 3.5 plants mP2did not differ significantly from that of the 6.3 plants n r 2 .
Vegetative Growth

Vertical Leaf Distribution Spatial leaf-area distribution for the 6.5 plants m-2 was more concentrated at the ear level ( ~ 0 . 9 m) when 0 compared to a more uniform vertical distribution for the low and medium PPD (Fig. 5). Dispersion of leaf area in space and time is an important factor in maximizing production of both sole and understory intercrops. The leaf area concentration around ear level corresponded to its greater light interception. It is consistent with earlier results (Loomis et al., 1968), which reported that the maximum leaf area was located at the ear stratum irrespective of PPD. The concentration of healthy leaf area and grain dry matter accumulation at the ear locus may explain the responsiveness of maize to increasing PPD. The ear leaves 'were longer and wider than others (data not shown), plus they were relatively young and had the shortest assimilate translocation pathway to the grain.

2'4b(0.1 below tassel 1

-0.2m

$
z a
U

0.9 +ear

level

4
"p
I
I
I

I (3
A

I-

20 -

a
0

1.9plants m-2 3.5 plants m-2 o 6.3 plants m-2

0.6

A 1.9 0 3.5

0.31 0 6.3 plants

I
1
I I I , I I I
I I
I !

.d 0

21

35

49

63

77

9 1

105

20

40

60

80
(/I "o

95100

LIGHT INTERCEPTION

DAYS AFTER PLANTING

Fig. 4 Influence by plant population density on light interception by maize canopy during grain filling in 1986. The broken horizontal line represents 95% interception, the critical leaf area index.

Fig. 3 Light interception profiles in maize canopy depths during grain filling as influenced by plant population density in 1986.

TETIO-KAGHO & GARDNER: MAIZE CANOPY STRUCTURE & LIGHT TRANSMISSION

933

Leaf Area Index Accumulation patterns of leaf area index (LAI) over time were similar for all three PPD (Fig. 6). Maxima LA1 of 1.7, 2.6, and 4.0 (63 DAP) were obtained for 1.9, 3.5, and 6.3 plants mP2,respectively, but LA1 declined thereafter. The LA1 maxima corresponded to nearly 95% or more light interception (critical LAI; Brougham, 1956) for the 3.5 and 6.3 plants m-2, but the 1.9 plants m-* intercepted less than 95%. The critical LA1 was 2.6 and 4.0 for the medium and high PPD, respectively. Pearce et al. (1 965) and Williams et al. (1968) reported a direct relationship between LA1 and light interception and resulting photosynthesis up to the critical LAI. In general, photosynthesis increases until nearly all incident solar radiation is intercepted by photosynthetic surfaces, and any further increase in leaf area only increases shading of the lower leaves with little benefit to the plant (Gardner et al., 1985).
24[ /(O.I-0.2m
2.1

Dry Matter Accumulation Plant population density significantly affected dry matter (DM) accumulation in the leaf, stem,and stalk (leaf plus stem) (Fig. 7), and the growth pattern for these different components was similar. Leaf and stem growth did not differ significantly from each other until 49 DAP; thereafter, the stem gradually accounted for more than 50% of the stalk DM. After about 70 DAP, stem and leaf, especially at the lowest PPD, gradually lost DM, probably by remobilization of stem solubles to grain. Leaf, stem, and stalk DM yield did not differ significantly among the three PPD until after 49 DAP, indicating the onset of interplant competition. The final result was 20 to 50% less yield per plant for each component from the 3.5 and 6.3 plants rn-

5r

1.9piants m-2

below tassel)

I .0

I
I -

I .5

A I .9 plants m-2 3.5plants m-2 0 6.3plants m-2

n w
0

I .2
09
0.6

a W a
L A

2 0
a v

a W

0.3
1 l l l l l , l , l , ,

6
0
1 4

LEAF AREANOLUME

(m2m-3)

28

42

56

70

84

98 105

Fig. 5 Influence of plant population density on vertical distribution of leaf area volume in maize canopy during the grain filling in 1986.

DAYS AFTER PLANTING Fig. 6 Influence of plant population density on leaf area index accumulation in maize, mean of 1985 and 1986.
p40

210

180 0 stalk

190
120

6.3 plants d 2
Oleaf

A stem

90
60

90
60

30
0

30
0

DAYS AFTER PLANTING Fig. 7 Leaf, stem and stalk (leaf plus stem) dry matter accumulation in maize at three plant population densities (1.9, 3.5, and 6.3 plants m-), mean of 1985 and 1986.

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AGRONOMY JOURNAL, VOL. 80, NOVEMBER-DECEMBER 1988

treatments, respectively, compared with the 1.9 plants mP2.

Crop Growth Rate Crop growth rate was significantly affected by PPD (Table 2). The 6.3 plants mF2accumulated 41.6 and 103.5% more dry matter per unit land area per day than the 3.5 and 1.9 plantsp2,respectively. The greater CGR obtained at the higher PPD corresponded closely to its greater LA1 and light interception.

duction in the lower canopy, probably inhibited sitem growth in the ultra-high PPD.

Plant Height Plant height increased to a maximum and then decreased (parabolically) with increasing PPD (Fig. 8), which was similar to the trend reported by Stinson and Moss (1960) and Early et al. (1966). Plant height was greatest at 6 to 10 plants mp2.Internode elongation (etiolation), due to shade effect, is believed to be an auxin response based on the theory that there is photodestruction of auxin at high irradiance, which results in reduced plant height (Leopole and Kriedemann, 1975). However, the decrease of plant height at ultra-high PPD was probably associated with limitations of assimilate and perhaps minerals and water. Competition for growth factors, added to the light reTable 2. Crop growth rate (CGR) influenced by plant populaas tion density (PPD) in maize, mean of 1985 and 1986.
PPD
plants m-'

Tillering The number of tillers per plant in the 1986 experiment decreased linearly as P'PD increased up to 3.5 plants mL2 (Fig. 9). Plants at greater PPD did not tiller. The formation of tillers appeared to be highly controlled by the environment. In 1985, few tillers were produced, even at low PPD, but in 1986 the higher irradiance in May and the larger amounts of N applied early in the season apparently promoted more tiller formation. However, tiller production in 1986, contrasted with 1985, was at the expense of the third ear, even at the low PPD (0.8-3.5 plants m-2).
CONCLUSIONS Increasing PPD in maize, up to 3.5 plants m - 2 or more, increased LA1 to the critical value (95% light interception). Also, increasing PPD increased LA1 concentration and light capture in upper canopy, particularly at the ear locus, perhaps an advantage in the source-sink relationship. Tassels intercepted from 2 to 40% solar radiation over the range of 0.8 to 15 plants m-*, but the assimilate balance related to this aspect was not assessed and needs more study. The canopy intercepted at least 40% irradiance at the low PPD (1.9 plants mp2) as early as 35 DAP, which suggests that an understory inter- or relay-crop (an interest in this study) will receive only 60% or less of incident light early in the season and much less than this amount later. Our data also show that light measurements in canopy do not have to be made at midday, as commonly believed, for reliable results; rather, they can

CGR
g m-' d-'

19 . 35 . 6.3 LSD (0.05)

14.2 20.4 28.9

3.4

3.5 3 0.
2.5

0 0

E 20. I-

1.5 -

y29 .3

1 0 . 9 - 0.68 ~~

x , R2=0.57 k

LL

00

0
[L

10 .
0.5 1

5 z

I -

\
2

2 . 9 - 0 . 7 6 ~ ~= 0 . 8 0 , r2

10-

12

1 4

1 6

0 0

'

IO

12

1 4

1 6

DENSITY (plants
Fig. 8 Relationship between maize plant height and plant population densities in 1986.

DENSITY (plants m-2) Fig. 9 Relationship between number of tillers per plant and imaize plant population density (restricted to only densities where the tillers were produced) in 1986.

TETIO-KAGHO & GARDNER: MAIZE REPRODUCTIVE DEVELOPMENT & YIELD

935

be made anytime after sunrise during clear morning hours. Morning solar radiation is the image of that in afternoon, but afternoon measurements are generally less successful due to more cloud formations.