1 Plant Population and Hybrid Impacts on Corn Grain and Forage Yield and 2 Nutrient Uptake 3 4 ABSTRACT 5 Corn (Zea mays L.) production recommendations must be periodically evaluated to 6 ensure that production practices remain in step with hybrid genetic improvements. Since 7 most of the recent increases in corn grain yield are due to increased optimum plant 8 density and not increased per plant yield, this study was undertaken to measure the 9 effects of plant population and hybrid on corn forage and grain yield and nutrient uptake. 10 Plant population (4.9, 6.2, 7.4, and 8.6 seeds m-2) and corn hybrid relative maturity (RM) 11 [early (108 day RM); medium (114 day RM); and late (118 day RM)] combinations were 12 evaluated in five site years under irrigated and non-irrigated conditions. The interaction 13 of hybrid with plant density was minimal. The latest RM hybrid outyielded the medium 14 and early hybrids by 550 and 1864 kg ha-1, respectively. Grain yield was highest at 8.5 15 plants m-2. Total stem yield was also greatest at the highest plant density but only 340 kg 16 ha-1 more than at 7.4 seeds m-2. Based on grain yield response over sites, the estimated 17 optimum population was 7.6 seeds m-2 which is 0.7 seeds m-2 higher than the current 18 recommendation at this average yield level (11.5 Mg ha-1). Grain nitrogen (N), 19 phosphorus (P), and potassium (K) uptake were highest for the medium hybrid and 20 resulted from greater nutrient concentrations and not yield. Nutrient uptake levels varied 21 by population with the lowest levels observed at the lowest and highest plant densities. 22 At 4.9 seeds m-2 this is explained by lower biomass yield. At the 8.6 seeds m-2 rate, N 23 and K supply may have been limiting resulting in lower overall concentrations. 1 1 INTRODUCTION 2 Plant Density 3 Recent increases in corn yields are attributed to greater stress tolerance of modern 4 hybrids, especially stress from interplant competition (Tokatlidis and Koutroubas, 2004). 5 Most of the increase in yield per unit area has been the result of increased optimum plant 6 population and not increased grain yield per plant. This is the result of more efficient 7 capture and use of resources such as water, sunlight and nutrients. Modern hybrids have 8 greater leaf longevity, more efficient root systems, and greater assimilate supply available 9 for translocation to developing grain (Tollenaar and Wu, 1999). 10 While relative corn grain yield levels have varied by environment, researchers 11 have recently reported maximum yields at or near the highest studied populations in a 12 number of locations (Nafziger, 1994; Staggenborg et al., 1999; Stanger and Lauer, 2006; 13 Thomison and Jordan, 1995; Widdicombe and Thelen, 2002) and not the parabolic 14 relationship previously reported (Alessi and Power, 1974; Karlen and Camp, 1985). On 15 sandy, coastal plain soils Karlen and Camp (1985) found that increasing plant density 16 from 7 to approximately 10 plants m-2 resulted in yield decreases unless supplemental 17 irrigation was applied. Recent research in the mid-Atlantic evaluating the effect of row 18 spacing and population found a significant interaction of these factors in only one of 28 19 instances (Kratochvil and Taylor, 2005). However under relatively high-yielding 20 environmental conditions (9.5 Mg ha-1), grain yields did not decrease in response to high 21 plant densities until the threshold of 10 plants m-2 was passed. 22 Total corn biomass, measured as silage yield, has been shown to be influenced by 23 plant density in New York, USA, where maximum economic forage yield was calculated 2 1 to occur at 9.8 plants m-2 (Cox et al., 1998). These authors found no interaction of row 2 spacing and plant population for forage yield. 3 South Carolina research reported that at a plant density of approximately 7 plants 4 m-2, leaf area index (LAI) averaged 4.3 and increased to 5.9 when population was 5 increased to 10 plants m-2 (Karlen and Camp, 1985). These authors concluded that there 6 was no net advantage to the higher LAI at the higher population since the threshold level 7 of 3.5 to 4.5 (Eik and Hanway, 1966) was reached with the lower population. 8 9 Harvest index (HI) is known to decrease with increased plant density, especially when plant density is above that required for maximum yield (Center and Camper, 1973; 10 DeLoughery and Crookston, 1979). Recent research has found this to be the result of 11 increasing barrenness at higher densities (Tollenaar et al., 2006). Kiniry and Echarte 12 (2005) report a general response of HI to plant density. Reviewing four recently 13 published studies, HI remained constant when plant density was less than 10 plants-2, but 14 above this threshold, HI decreased at a rate of -0.012 units per plants m-2. 15 Nutrient concentration is generally held to be independent on plant density 16 whether for grain (Ottman and Welch, 1989; Overman et al., 2006) or forage (Hoff and 17 Mederski, 1960). However increasing plant density through narrower rows has decreased 18 grain N concentration (Stickler, 1964). In instances where plant populations are 19 insufficient to use all applied nitrogen, N concentration in grain and forage rises with 20 increased N rates, but at appropriate populations, higher N rates were used effectively 21 (Jordan et al., 1950) 22 Hybrids 3 1 In a corn forage study, Cox et al. (1998) studied eight hybrids with a range in RM 2 of 100 to 112 days across a range of plant populations and found that while there were 3 yield differences among hybrids, interactions with population were uncommon. 4 Deloughery and Crookston (1979) report that HI decreased with RM (75 day to 130 days 5 for ten hybrids) but only with environmental stress. 6 Nutrient concentration is known to vary widely among corn hybrids and is highly 7 dependent on environment (Ferguson et al., 1991; Heckman et al., 2003; Schenk and 8 Barber, 1980). Average reported values corn grain and 12.9, 3.8, and 4.8 g kg-1 of N, P, 9 and K, respectively (Heckman et al., 2003). 10 In row spacing by plant density studies in Indiana, hybrid RM differences had 11 little effect on grain yield, but greater stalk breakage was noted for one hybrid (Neilsen, 12 1988). Nafziger (1994) also found no hybrid by plant density interactions for two 13 hybrids. In an eastern Corn Belt study of plant population and corn hybrid prolificacy it 14 was concluded that environment, genetics, and plant population main effects were more 15 important to optimum grain yield than the hybrid by population interaction (Thomison 16 and Jordan, 1995). In eastern Nebraska, a later RM hybrid was found to produce greater 17 forage yield while the earlier hybrid had higher grain yield (Alessi and Power, 1974)). 18 Neither forage or grain yield by RM interacted significantly with plant density. 19 Conversely, recent Michigan research found significant interactions for plant density and 20 hybrid for grain yield and moisture of six hybrids (Widdicombe and Thelen, 2002). In 21 spite of the fact that the majority of published studies report no, or occasional, RM by 22 plant populations interactions, many growers and practitioners believe this to be the case. 4 1 The objectives of this research were to examine the effect of plant density and corn 2 hybrids of different RM on corn biomass and grain yields and nutrient uptake. 3 4 5 MATERIALS AND METHODS Small plot field studies were conducted in 2005 near Blacksburg, VA (37 12’ N, 6 80 34’ W) on a Hayter loam (fine loamy, mixed, active, mesic Ultic Hapludalf) and Mt 7 Holly, VA (38 5’ N, 76 43’ W) on a State fine sandy loam (fine loamy, mixed, 8 semiactive, thermic, Typic Hapludalf) and in 2006 at Mt. Holly. The Blacksburg site was 9 non-irrigated while trials were conducted under both irrigated and non-irrigated 10 conditions at Mt. Holly. Experimental design was a randomized complete block with a 11 split plot arrangement of treatments and four replications. Main plots were plant 12 population (4.9, 6.2, 7.4, and 8.6 seeds m-2) and subplots were corn hybrid relative 13 maturity [early – Pioneer® Brand ‘34B97’ (108 day RM); medium – Pioneer® Brand 14 ‘33M54’ (114 day RM); and late – Pioneer® Brand ‘31G66’ (118 day RM)]. Planting 15 dates and agronomic information for the five site years are listed in Table 1. Plots were 16 planted with a Wintersteiger 2600 vacuum plot planter (Wintersteiger Inc., Salt Lake 17 City, UT) and were four, 76-cm rows wide by 8 m long. In all cases the previous crop 18 was soybean at Mt. Holly trials and corn at the Blacksburg site. Starter fertilizer at a rate 19 of 43 kg N ha-1 and 5 kg P ha-1 was applied 5 cm below and 5 cm to the side of the seed 20 at planting. Total N rates in 2005 were 263, 190, and 170 kg N ha-1 at Mt. Holly 21 irrigated, Mt. Holly non-irrigated, and Blacksburg, respectively. Total N applied in 2006 22 was 252 kg ha-1 at the irrigated Mt. Holly site and 180 kg N ha-1 for the non-irrigated site. 23 Phosphorus and K were broadcast prior to planting at rates indicated by Virginia Tech 5 1 soil test recommendations (Donohue and Heckendorn, 1994). Planting, emergence, and 2 harvest dates as well as weather information were collected at all locations (Table 1). 3 Weather information was gathered using Watchdog™ (Spectrum Technologies Inc. 4 Plainfield, Illinois) weather stations at each site. 5 Aboveground biomass was hand harvested from each plot at the R6 6 developmental stages (Ritchie et al., 1992). A total of five consecutive plants in the outer 7 two rows were harvested at soil level. Total plant biomass as well as stem, leaf, and 8 reproductive biomass dry matter were determined by separating the components and 9 drying them to a constant weight in a forced-air oven at 60° C. Biomass yield (kg ha-1) 10 was calculated as the product of individual plant weight and measured plant population. 11 Stem and leaf tissue and grain from the R6 sampling date were ground to pass a 1- 12 mm screen. Total carbon and N concentration for each sample was determined via a 13 LECO Tru-Spec® CHN dry combustion analyzer (LECO Corporation, St. Joseph, MI). 14 Samples were digested using HNO3/HClO4 acid solution. Phosphorus (P) concentration 15 was determined colorimetrically (Kuo, 1996) and K concentration determined by atomic 16 absorption spectroscopy (Helmke and Sparks, 1996). Nitrogen, P, and K uptake for 17 grain and forage (stem and leaf ) was calculated as the product of biomass yield of the 18 respective plant component and nutrient concentration (%) within each component. 19 Grain was harvested from the center two rows from each plot using a Massey 20 Ferguson 8XP plot combine. Plot weight, grain moisture, and test weight were 21 determined using a Graingage™ system (Juniper Systems, Logan, UT). Grain yield from 22 all trials is reported at 155 g kg-1 moisture. Harvest index (HI) was calculated by 23 dividing grain yield by total plant biomass. 6 1 2 Statistical analysis was performed using the GLM procedure available from SAS 3 (SAS Inst., 2004). Due to interaction effects of treatments across years and locations, 4 data from each site year were analyzed and presented separately. Mean comparisons 5 using a protected LSD test were made to separate RM and plant population effects where 6 F-tests indicated that significant differences existed (P<0.10). Regression was applied to 7 evaluate plant density impacts on grain and forage yield and nutrient uptake across sites. 8 RESULTS AND DISCUSSION 9 Blacksburg 2005 10 Leaf yield was influenced by both plant density and corn hybrid (Table 2). As 11 population increased from 4.9 to 8.6 seeds m-2 leaf weight increased in a linear manner 12 (526 kg ha-1) but then decreased at the highest population. The medium maturity hybrid 13 had the greatest final leaf weight. Grain yield was more than 2600 kg ha-1 lower for the 14 early maturity hybrid than for the other two entries (Table 2). This is a river bottom site 15 typified by foggy, hazy mornings and relative cool nights. Because of the relatively 16 cooler night time temperatures, longer season hybrids typically offer a yield advantage. 17 Similar to the trends observed for leaf yield, increasing plant population up to 7.4 seeds 18 m-2 resulted in increased grain N and P uptake but with the further increase in plant 19 population to 8.6 seeds m-2 a decrease in uptake of 51 kg N and 12 kg P ha-1 was 20 observed (Table 2). Leaf+stem K uptake was higher at 7.4 seeds m-2 than at any other 21 population. 22 Mt. Holly Non-Irrigated 2005 23 Total stem, leaf, and grain yield was significantly lower for the early season 24 hybrid at this site (Table 3). Later maturing hybrids often accumulate more leaf and 7 1 stem biomass and thus more nutrients and carbohydrates that can be translocated to the 2 grain after anthesis. This allows more total assimilate to be available for grain production 3 in these longer season hybrids. Total grain N uptake was 31 kg ha-1 greater for the 4 medium maturity than for the early hybrid, again similar to the results observed for plant 5 growth parameters (Table 3). There was a significant interaction of population and 6 relative maturity for leaf+stem N uptake at this site, however main effects are still 7 presented for the sake of consistency. This was the only instance that an interaction 8 occurred for any of the investigated variables and in this case was due to low uptake 9 levels associated with the lowest population. 10 Mt Holly Irrigated 2005 11 Stem yield again exhibited a quadratic response to plant population with the 12 highest yield of 11,482 kg ha-1 reached at a population of 7.4 seeds m-2 (Table 4). The 13 early hybrid had approximately 700 kg ha-1 less stem dry weight than the medium or late 14 hybrids. Total leaf dry matter was higher at the two higher population treatments 15 compared to the lower populations (average of 3611 versus 2689 kg ha-1) (Table 4). Leaf 16 mass was again lowest for the early hybrid. Grain yield was lowest (13,713 kg ha-1) for 17 the lowest population and was increased by 1,400 kg ha-1 by increasing population to 6.2 18 seeds m-2. Grain P and K uptake were also lowest for the lowest population density at 19 this site (Table 4). 20 Mt Holly Non-Irrigated 2006 Stem yield was lower at 6.2 seeds m-2 (6150 kg ha-1) than at the 4.9 or 8.6 seed m- 21 22 2 rates but was not different from 7.4 seeds m-2 (Table 5). The late maturity hybrid 23 produced greater stem biomass yield than either of the others, probably due to the 8 1 expected longer vegetative growth period. A linear increase of leaf yield in response to 2 plant density was observed at this site. Leaf yield increased at a rate of nearly 300 kg ha-1 3 for each additional plant m-2 (Table 5). A similar trend was observed in response to 4 increasing relative maturity with the greatest leaf yield of 2961 kg ha-1 associated with 5 the late hybrid. This was the only site year in which HI was impacted by treatment. 6 Harvest index was significantly lower at 4.9 seeds m-2 than at other densities, probably 7 due to the high relative stem yield (8135 kg ha-1) in this treatment (Table 5). The late 8 hybrid had the lowest HI of 0.301. Significant differences are the result of greater leaf 9 yield at similar a grain yield. Grain P and K uptake were lowest at 6.2 seeds m-2 and 10 11 12 highest at the 7.4 seeds m-2 density. Mt Holly Irrigated 2006 Stem yield was lower for the medium RM hybrid compared to the late hybrid but 13 was similar to the early hybrid (Table 6). Leaf yield was 874 kg ha-1 greater at the 8.6 14 seeds m-2 density than at 4.9 seeds m-2, but no differences were observed among the other 15 treatments. Leaf biomass yield increased by 429 kg ha-1 for the medium versus the early 16 hybrid and 480 kg ha-1 for the late versus the medium hybrid (Table 6). Grain N uptake 17 was significantly lower at 8.6 seeds m-2 which might indicate a deficiency at this high 18 plant density. The lowest observed grain N uptake (135 kg ha-1) was associated with the 19 late maturity hybrid while there were no other differences by maturity. Conversely, 20 stem+leaf N uptake was highest in the late hybrid and lowest (59 kg ha-1) for the medium 21 hybrid. Grain P uptake was least at 4.9 seeds m-2 and this corresponds to the numerically 22 lowest grain yield. The late RM hybrid also had slightly lower grain P uptake than the 23 others. Grain K uptake was lowest at 4.9 seeds m-2 (35 kg K ha-1) and again is explained 9 1 mostly by the lower numerical yield for this treatment. Similar grain K uptake was noted 2 between the early and late hybrids with significantly more (4 kg ha-1) K taken up by the 3 medium hybrid. 4 5 Plant population effects on yield and nutrient uptake Overall, grain yield increased in a quadratic manner with increasing plant density 6 (Figure 1) and while the rate of increase was declining, the yield was greatest at the 7 highest plant density. The impact of plant density in these studies is similar to that 8 reported by Kratochvil and Taylor (2005) and Widdicombe and Thelen (2002) 9 (approximately 300 kg ha-1 increase for each seed m-2) and more than that reported by 10 others (Farnham, 2001; Karlen and Camp, 1985). This may be due to the ability of the 11 specific hybrids chosen to compensate with larger ears in response to less interplant 12 competition. The growing environment was also favorable in all years of this study and 13 so plant to plant competition for water may have been less a factor than would be seen in 14 many instances. Stem and leaf yield also generally increased with increasing plant 15 density (Figure 1). Grain N uptake increased incrementally up to 7.4 seeds m-2 but 16 decreased in the last increment (Figure 2). This same trend was evident in forage N and 17 K uptake (Figure 3) and may be the result of nutrient limitations at these high plant 18 densities since yield was still increased. Grain P and K uptake were similar across the 19 range of densities at 38 and 42 kg ha-1, respectively. Leaf+stem P uptake was also 20 essentially the same (17 kg P ha-1) across plant densities (Figure 3). 21 22 23 CONCLUSIONS Grain yield was highest at the highest plant density with a total advantage of 1141 kg ha-1 over the 4.9 seeds m-2 treatment. The late hybrid outyielded the medium 10 1 and early hybrids by 550 and 1864 kg ha-1, respectively. Total stem yield was also 2 greatest at the highest plant density but only 340 kg ha-1 more than at 7.4 seeds m-2. 3 Overall results for leaf yields were similar to those for stem yield. Solving for the 4 optimum population based on the quadratic model for grain yield results in an estimated 5 optimum of 7.6 seeds m-2. This is slightly higher than the current recommendation of 6.9 6 seeds m-2 that would be recommended for this average yield level (11.5 Mg ha-1). 7 Harvest index was generally unaffected by treatment in these studies. This is not 8 surprising given the findings of Kiniry et al. (2005) who report a basically constant HI 9 with plant densities below 10 plants m-2, at which point HI begins to decrease at a rate of 10 11 -0.012 units per plant m-2 increase. Grain N, P, and K uptake was highest for the medium RM hybrid. Since grain 12 yield for this hybrid was not the highest, these uptake values result from greater nutrient 13 concentrations in the grain. Leaf+stem N uptake for this medium hybrid was the lowest 14 of the three but K uptake was highest. Nutrient uptake levels varied somewhat by 15 population with the lowest levels observed at the lowest and highest plant densities. At 16 4.9 seeds m-2 this is explained by generally lower biomass yield. At the 8.6 seeds m-2 17 rate, N and K may have been limiting resulting in lower overall concentrations in tissue. 18 While numerous studies over the years have investigated the effect of plant 19 population and hybrid characteristics on yield, recent increases in corn hybrid stress 20 tolerance and standability make revisiting these basic studies worthwhile. Based on 21 results from these studies, plant density recommendations at these high yield levels may 22 need to be revised upwards. Correspondingly, recommended N and K nutrient levels 23 should be evaluated to ensure that adequate but not excessive supplies are available. To 11 1 meet the needs of plant densities this high, fertilizer rates may well need to increase 2 concomitantly. 12 Table 1. Production practices, sampling dates, and environmental conditions, 2005-2006. Total N applied ---kg ha-1--263 R6 Biomass Sampling Date ---date--09/01/2005 GDD Planting to Sampling ----°C---1688 Rainfall Planting to Sampling ----mm---490 30 yr-mean growing season rainfall ----mm---455 Grain Harvest ---date--09/20/2005 04/20/2005 190 09/01/2005 1689 493 455 09/19/2005 05/09/2005 05/14/2005 170 09/20/2005 1484 394 452 10/05/2005 MT Holly Irrigated 04/26/2006 05/03/2006 252 08/30/2006 1665 410 456 09/20/2006 MT Holly Non-irrigated 04/28/2006 05/04/2006 180 08/29/2006 1658 409 456 09/22/2006 MT Holly Irrigated Planting Date ---date--04/19/2005 Emergence Date ---date--04/25/2005 MT Holly Non-irrigated 04/13/2005 Blacksburg Experiment 13 Table 2. Analysis of variance and means for stem, leaf and grain yield, harvest index, and N, P and K uptake in response to plant population and relative maturity, Blacksburg, 2005 Source of Variation Block Population Block*Population Relative Maturity (RM) Population*RM Block*Population*RM Density ---plants m-2--4.9 6.2 7.4 8.6 LSD RM Early Medium Late LSD df 3 3 9 2 6 24 Stem Yield 2095962 15874801 13019592 4145858 2434027 30950870 Leaf Yield 268893 1691979 *** 383085 952804 *** 301302 1133168 Grain Yield 2970054 17742849 7037293 15378885 * 3838368 12614450 -----------------------------kg ha-1----------------------------4281 855 11129 4678 1061 11593 5845 1381 12295 4899 1126 12018 ns 191 ns 4525 5030 5222 ns 969 1300 1048 163 9776 12399 13103 1616 HI 0.011 0.009 0.008 0.019 0.005 0.013 ----------N uptake---------Grain Leaf+Stem 2297 106 4572 * 44 840 27 1084 32 462 143 4206 56 ---------P uptake---------Grain Leaf+Stem 90 5 209 * 2 38 3 34 1 31 9 324 5 ----------K uptake---------Grain Leaf+Stem 29 4 25 1942 ** 133 114 147 1337 46 454 236 1059 --------------------------------------------------------kg ha-1-------------------------------------------------------0.478 170 46 36 10 36 105 0.513 177 45 37 9 37 110 0.526 181 51 41 10 41 144 0.470 130 45 29 10 47 111 ns 43 ns 11 ns ns 20 0.535 0.469 0.486 ns ***,**,* - Significant at the 0.01, 0.05, and 0.10 level, respectively. 14 149 173 171 ns 44 48 48 ns 32 39 37 ns 10 10 9 ns 32 45 44 ns 111 132 109 ns Table 3. Analysis of variance and means for stem, leaf and grain yield, harvest index, and N, P and K uptake in response to plant population and relative maturity, Mt Holly Non-Irrigated, 2005 Source of Variation Block Population Block*Population Relative Maturity (RM) Population*RM Block*Population*RM Density ---plants m-2--4.9 6.2 7.4 8.6 LSD RM Early Medium Late LSD df 3 3 9 2 6 24 Stem Yield 27530833 30619921 40339801 35396227 ** 22674926 89683643 Grain Yield 32016330 95105621 *** 11032462 54997876 ** 8104451 25376120 HI 0.011 0.005 0.008 0.001 0.004 0.030 -----------------------------kg ha-1----------------------------6339 1675 11988 7903 2161 12451 8452 2637 13123 8008 2302 13909 ns 601 1868 0.469 0.439 0.490 0.466 ns 6463 8342 8222 1388 Leaf Yield 2870702 5741041 ** 3817424 5085059 *** 1866044 5760509 1734 2401 2446 363 11321 13643 13639 2266 0.472 0.459 0.468 ns ***,**,* - Significant at the 0.01, 0.05, and 0.10 level, respectively. 15 ----------N uptake---------Grain Leaf+Stem 3489 6 9547 1952 3781 1105 4279 * 25 1383 893 * 2966 709 ---------P uptake---------Grain Leaf+Stem 53 61 548 27 172 72 141 30 73 71 144 56 ----------K uptake---------Grain Leaf+Stem 63 158 527 8720 179 3336 144 3486 49 1396 127 3021 --------------------------------------------------------kg ha-1-------------------------------------------------------171 57 42 20 44 126 193 99 46 24 46 210 185 88 46 21 46 203 198 75 50 23 50 182 ns ns ns ns ns ns 173 204 183 23 79 78 82 ns 43 47 48 ns 23 23 20 ns 43 49 48 ns 163 204 174 ns Table 4. Analysis of variance and means for stem, leaf and grain yield, harvest index, and N, P and K uptake in response to plant population and relative maturity, Mt Holly Irrigated, 2005 Source of Variation Block Population Block*Population Relative Maturity (RM) Population*RM Block*Population*RM Density ---plants m-2--4.9 6.2 7.4 8.6 LSD RM Early Medium Late LSD df 3 3 9 2 6 24 Stem Yield 6805014 66458345 ** 33769250 17248025 ** 2522255 52843379 Grain Yield 4552003 167512818 *** 4133849 21092372 ** 7028214 7725426 HI 0.005 0.013 0.005 0.006 0.002 0.010 -----------------------------kg ha-1----------------------------8842 2614 13713 8791 2763 15149 11482 3692 14887 10738 3530 15090 1789 322 1378 0.411 0.422 0.470 0.476 ns 9241 9939 10709 1142 Leaf Yield 498847 10503882 *** 1093256 5706049 *** 1031401 3772209 2699 3216 3536 329 13267 14727 16134 1733 0.464 0.446 0.424 ns ***,**,* - Significant at the 0.01, 0.05, and 0.10 level, respectively. 16 ----------N uptake---------Grain Leaf+Stem 613 857 10586 351 514 2935 2169 176 1347 929 1401 1511 ---------P uptake---------Grain Leaf+Stem 28 216 606 ** 77 30 532 64 41 73 86 114 456 ----------K uptake---------Grain Leaf+Stem 27 12843 678 ** 10498 62 9367 95 7239 86 5928 111 9420 --------------------------------------------------------kg ha-1-------------------------------------------------------220 130 46 28 48 215 242 137 53 29 55 235 234 149 52 31 54 281 232 137 53 27 55 242 ns ns 8 ns 6 ns 214 277 206 ns 144 137 135 ns 47 55 51 ns 29 31 26 ns 47 58 53 ns 236 266 228 ns Table 5. Analysis of variance and means for stem, leaf and grain yield, harvest index, and N, P and K uptake in response to plant population and relative maturity, Mt Holly Non-Irrigated, 2006. Source of Variation Block Population Block*Population Relative Maturity (RM) Population*RM Block*Population*RM Density ---plants m-2--4.9 6.2 7.4 8.6 LSD RM Early Medium Late LSD df 3 3 9 2 6 24 Stem Yield 7058667.63 26681189 * 28564724.6 31564665.7 ** 17247575.8 41169416.6 Leaf Yield 498633.92 9599699.4 *** 1531544.4 5358128 *** 1143356.7 3460839.7 Grain Yield 15664242 5275737 7097692 1642716 1424773 3901866 HI 0.004 0.003 ** 0.001 0.006 * 0.002 0.006 -----------------------------kg ha-1----------------------------8135 2190 7794 6150 2203 7001 7329 2579 8143 7818 3332 8025 1645 392 ns 6891 6676 8551 1044 2197 2547 2961 305 0.303 0.337 0.328 0.327 0.031 7517 7839 7867 ns 0.337 0.333 0.301 0.033 ***,**,* - Significant at the 0.01, 0.05, and 0.10 level, respectively. 17 ----------N uptake---------Grain Leaf+Stem 2452 257 1687 5598 423 1256 524 1484 683 1898 1034 2909 ---------P uptake---------Grain Leaf+Stem 78 52 39 ** 113 4 115 12 14 18 15 48 138 ----------K uptake---------Grain Leaf+Stem 62 5218 48 ** 2941 2 4376 4 532 18 8823 50 7253 --------------------------------------------------------kg ha-1-------------------------------------------------------99 50 24 12 32 154 88 38 21 7 29 148 103 44 25 11 34 175 109 41 25 11 31 188 ns ns 4 ns 2 ns 101 98 100 ns 44 36 52 ns 24 23 24 ns 11 9 10 ns 32 33 30 ns 160 169 164 ns Table 6. Analysis of variance and means for stem, leaf and grain yield, harvest index, and N, P and K uptake in response to plant population and relative maturity, Mt Holly Irrigated, 2006. Source of Variation Block Population Block*Population Relative Maturity (RM) Population*RM Block*Population*RM Density ---plants m-2--4.9 6.2 7.4 8.6 LSD RM Early Medium Late LSD df 3 3 9 2 6 24 Stem Yield 14625089.1 3476418.7 13461379.4 22842546.9 ** 7908444.35 41476981.2 Leaf Yield 2462992.1 4651133.7 ** 3207713.8 6613854.5 *** 1261407.7 4861178.9 Grain Yield 155664242 5275737 7097692 1642716 1424773 3901866 HI 0.007 0.007 0.002 0.013 0.005 0.007 -----------------------------kg ha-1----------------------------7295 2558 9754 7377 3022 11256 7487 3091 10664 7838 3432 11041 ns 551 ns 7208 6834 8506 1301 2580 3009 3489 316 0.317 0.321 0.309 0.269 ns 10577 10422 11038 ns 0.320 0.322 0.271 ns ***,**,* - Significant at the 0.01, 0.05, and 0.10 level, respectively. 18 ----------N uptake---------Grain Leaf+Stem 23 5 415 ** 449 31 97 35 826 ** 230 ** 374 70 240 ---------P uptake---------Grain Leaf+Stem 1 25 32 * 99 4 33 1 119 ** 9* 31 4 45 ----------K uptake---------Grain Leaf+Stem 1 22 55 ** 873 5 1321 23 * 150 11 3044 13 3051 --------------------------------------------------------kg ha-1-------------------------------------------------------150 95 29 17 35 197 151 52 36 14 43 162 154 87 35 14 41 196 138 55 34 12 45 180 10 ns 4 ns 4 ns 156 154 135 9 74 59 84 15 34 34 32 2 14 13 15 3 39 44 40 3 175 176 201 ns Figure 1. Average grain, stem, and leaf yield as affected by plant density. 14000 y = -68.54x2 + 1235.3x + 6452.7 R2 = 0.9987 Yield, kg ha -1 12000 10000 y = 55.462x2 - 496.16x + 8039.1 R2 = 0.9512 8000 Stem 6000 4000 Leaf y = -31.854x2 + 653.85x - 501.16 Grain R2 = 0.9632 2000 0 0 1 2 3 4 5 6 Plant Density, seeds m 19 7 -2 8 9 10 Figure 2. Grain N, P, and K uptake as affected by plant density. Grain Nutrient Uptake, kg ha-1 200 180 160 N 140 P y = -2.9491x2 + 40.011x + 36.176 120 K R2 = 0.9885 100 80 y = -0.781x2 + 11.36x - 1.671 60 R2 = 0.9972 40 2 y = -0.1012x + 3.102x + 26.226 R2 = 0.9883 20 0 0 1 2 3 4 5 6 7 -2 Plant Density, plants m 20 8 9 10 Leaf+stem Nutrient Uptake, kg ha-1 Figure 3. Leaf+stem N, P, and K uptake as affected by plant density. 250 y = -5.3935x2 + 80.539x - 109.72 R2 = 0.7944 200 N P 150 K y = -1.9127x2 + 25.608x - 5.9771 100 R2 = 0.3843 50 y = 0.002x2 - 0.2181x + 18.334 R2 = 0.3628 0 0 1 2 3 4 5 6 Plant Density, plants m 21 7 -2 8 9 10 REFERENCES Alessi, J., and J.F. 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