Crop Science 42:833-841 (2002)
© 2002 Crop Science Society of America
TURFGRASS SCIENCE
Fate and Transport of Nitrogen Applied to Six Warm-Season Turfgrasses
D. C. Bowman*,
C. T. Cherney and
T. W. Rufty, Jr.
Dep. of Crop Science, North Carolina State Univ., Raleigh, NC 27695
* Corresponding author (dan_bowman{at}ncsu.edu)
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ABSTRACT
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A greenhouse study compared six warm season turfgrasses {common bermudagrass [Cynodon dactylon (L.) Pers.], Tifway hybrid bermudagrass (C. dactylon x transvaalensis), centipedegrass (Eremochloa ophiuroides (Munro) Hack.), Raleigh St. Augustinegrass [Stenotaphrum secundatum (Walter) Kuntze], Meyer zoysiagrass (Zoysia japonica Steud.), and Emerald zoysiagrass (Z. japonica x tenuifolia)] for NO3-N leaching and N use efficiency. Sod was established in sand-filled columns and managed under worst-case conditions to promote nitrate leaching. Ammonium nitrate was applied at 50 kg N ha-1 on seven dates, with the final application labeled with 15N. Leachate samples were collected and analyzed for NO3-N and NH4-N and clippings were analyzed for total N. Leaching losses were high following the first N application, ranging from 48 to 100% of the NO3-N and 4 to 16% of the NH4-N applied. Nitrate loss from subsequent applications was reduced substantially, while NH4 leaching was essentially eliminated. There were significant differences among species for leachate NO3-N concentration and cumulative N leached, with St. Augustinegrass being the most effective and Meyer zoysiagrass the least effective at minimizing NO3 leaching. Nitrogen recovery by the turf ranged from 63% for Meyer zoysiagrass to 84% for hybrid bermudagrass. Root length density (RLD) varied significantly among species at depths >30 cm, and was negatively correlated with NO3 leaching loss. These results document differences between the warm season turfgrasses for NO3 leaching potential, possibly related to root distribution, and emphasize that species selection is an important factor in minimizing environmental impacts from turfgrass management.
Abbreviations: RLD, root length density
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INTRODUCTION
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NITROGEN IS THE MINERAL nutrient required in the greatest quantity by turfgrasses. When maintained at adequate levels, N promotes vigor, visual quality, recovery from damage, and overall health. The amount of available N in most soils is insufficient to support high quality turf, and regular applications of N fertilizer are thus required.
Nitrogen is a dynamic nutrient and undergoes numerous transformations and movement within the turf system (Petrovic, 1990). It enters the system primarily as fertilizer, and exits via gaseous loss, leaching, clipping removal, and under some conditions, surface runoff. Of these processes, the public perceives NO3 leaching as the greatest environmental threat.
Research conducted during the last three decades generally documents low potential for NO3 leaching from properly managed turf (Rieke and Ellis, 1974; Starr and DeRoo, 1981; Sheard et al., 1985; Gold et al., 1990; Mancino and Troll, 1990; Miltner et al., 1996). However, higher leaching losses have been associated with several factors, including irrigation management, turfgrass establishment, and species or cultivar selection.
Irrigation in excess of evapotranspirational demand, with the resulting downward movement of water, increased NO3 leaching from both Kentucky bluegrass (Morton et al., 1988) and hybrid bermudagrass (Snyder et al., 1984). In the latter study, daily irrigation resulted in NO3 leaching losses ranging from 22 to 56% of the applied N. By contrast, losses were reduced to <1% by scheduling irrigation based on soil moisture depletion, as determined with tensiometers. Nitrate leaching is also affected by the timing of irrigation relative to fertilizer application. Delaying irrigation for 5 d following N application reduced leaching losses up to 90% compared with turf irrigated 1 d after N application (Bowman et al., 1998).
Significant leaching may occur during turfgrass establishment. Geron et al. (1993) reported relatively high NO3 concentrations, averaging 14 mg N L-1, in leachate from a Kentucky bluegrass turf during the first year after planting. However, values decreased to
2 mg N L-1 during Year 2, indicative of a more mature turfgrass. They also noted that in Year 2, leachate NO3 concentrations were consistently lower from a seeded turf compared with a sodded turf, with volume weighted averages of 1.1 and 3.5 mg N L-1, respectively. The authors attributed the difference to more extensive root development in the seeded bluegrass.
Turfgrass genotype has been shown to affect NO3 leaching for several cool-season species. Liu et al. (1997) found both inter- and intraspecific differences for NO3 leaching potential in Kentucky bluegrass, perennial ryegrass, and tall fescue, with the ranking for NO3 loss being Kentucky bluegrass > perennial ryegrass > tall fescue. The authors speculated that these genotypic differences in leaching might be due to differences in NO3 uptake efficiency. Bowman et al. (1998) compared two genotypes of creeping bentgrass that differed in root density, and found that a more extensive root system reduced leaching losses by roughly 50%.
Most of the research on NO3 leaching from turfgrasses has focused on cool-season species (see above) or bermudagrass (Brown et al., 1977, 1982; Snyder et al., 1981). There are little or no data on NO3 leaching from the numerous other warm-season turfgrasses, in spite of their wide distribution and use. To address this deficiency, this study compared several of the common warm-season turfgrasses for NO3 leaching, N absorption, and N use efficiency.
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MATERIALS AND METHODS
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General Procedures
An experiment was conducted at the North Carolina State University Research Greenhouse facility (Raleigh, NC) beginning in July 1996 and ending October 1997. Average maximum and minimum temperatures were
30 and 21°C, respectively, during fall through spring, and 34 and 22°C, respectively, during the summer.
Six warm-season turfgrasses (common bermudagrass, Tifway hybrid bermudagrass, centipedegrass, Raleigh St. Augustinegrass, Meyer zoysiagrass, and Emerald zoysiagrass) were established in polyvinyl chloride column lysimeters, 35 cm in diameter and 75 cm deep. Three porous ceramic extraction cups, joined with a cross connector, were placed at the bottom of each column. The cups were connected to 3-L collection bottles, which were connected to a vacuum manifold. A 7-cm layer of diatomaceous earth covered the ceramic cups to prevent their plugging. The columns were then filled with a coarse sand (Table 1) to a final bulk density of 1.6 g cm-3. A preplant fertilizer supplying 50 kg N ha-1, 100 kg P ha-1, and 100 kg K ha-1 was incorporated to a depth of 4 cm. Commercially grown sod of each species was washed free of soil, cut to fit the lysimeters, and planted on 18 July, 1996. The sod was lightly irrigated twice daily during the first 4 wk following planting. Orthene was applied 20 Dec., 25 Apr., and 29 May to control mealybugs and aphids.
Fertilizer Treatments
Nitrogen was applied as NH4NO3 (34-0-0) at a rate of 50 kg N ha-1 on 23 Oct. and 22 Nov. 1996, and on 21 Mar., 6 May, 7 June, 18 July, and 15 Sept. 1997. The N source for the September application was 15NH154NO3, with an enrichment of 10.0%. Fertilizer was applied as a granular material and watered in with 1 cm of irrigation.
Phosphorus and K were each applied at 50 kg ha-1 using triple super phosphate (0-46-0) and KCl (0-0-60) on 10 Oct. 1996, and 14 Jan., 14 Mar., and 10 Sept. 1997. Micronutrients (Micromixâ, Lesco Inc., Strongsville, OH) were applied at the recommended rate four times during the study. Foliar applications of FeSO4·7H2O (1.8 kg Fe ha-1) were made three times during the experiment to prevent iron deficiency.
Leachate Sampling
Evapotranspiration was estimated using gravimetric water loss from mini-pan lysimeters positioned at canopy height. Columns were irrigated three times per week to provide a target leaching fraction of 50%, based on estimated evapotranspiration (total weekly irrigation ranged from 410 cm of water). Leachate was collected following each irrigation by applying a tension of 0.02 MPa to the columns for 15 h. Leachate volume was recorded and a subsample collected for analysis. Nitrate and NH+4 in the leachate were determined by the rapid diffusion method (Carlson, 1986). Volume weighted concentrations were calculated as the total amount of NO3 or NH4 leached (mg N column-1) divided by the total leachate volume (L column-1) for a given sampling period.
Turf cultures were mowed regularly at 5 cm and the clippings were collected, oven dried, weighed, and ground. Tissue N was determined by Kjeldahl digestion. Root length density was determined at the conclusion of the study. A 5-cm diameter soil core was extracted from the center of each column, and subsamples were cut from each core at depths of 4 to 6, 17 to 19, 29 to 31, 42 to 44, 54 to 56, and 67 to 69 cm. Roots were separated from soil by elutriation and quantified using the modified line intersect method (Tennant, 1975). Shoot tissue was excised at the soil surface, roots were separated from the sand, and both tissues were processed as for the clippings. All tissues were then analyzed for total N and 15N enrichment by mass spectrometry.
The experiment was conducted as a randomized complete block design with six treatments (species) and four replicates. Data were analyzed by ANOVA and means separated at P = 0.05 by the least significant difference test (SAS Institute, 1995).
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RESULTS AND DISCUSSION
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The six warm-season turfgrasses used in this study are grown throughout the southeastern USA for home lawns, golf courses, athletic fields, right-of-ways, and various other applications. They exhibit a wide range of fertility requirements, growth rates, and water use. Bermudagrass, for example, requires relatively high N rates, whereas centipedegrass thrives on little to no N, and may even deteriorate with higher N inputs (Duble, 1996). The design of this study did not permit consideration of individual fertility requirements, since all species were treated identically.
Experimental conditions were chosen to increase the potential for NO3 leaching. A highly porous sand was used as the root zone, fertility was maintained at a fairly high level (350 kg N ha-1 yr-1), and irrigation was scheduled to provide a 50% leaching fraction. The conditions are realistic for recently sodded or sprigged warm season turfgrass growing in the southern portion of their range, but would be somewhat excessive for a mature stand.
Sod establishment was slow due to the low water holding capacity of the rootzone and elevated temperatures in the greenhouse. It took
3 mo for the turfgrasses to root sufficiently such that they could not be pulled up by gently tugging on the sod. Although a complete fertilizer was applied prior to planting, leaching data were not collected during the first 3 mo, due to the lack of significant rooting. On the basis of subsequent data, it is likely that much of the pre-plant N leached. Once the sod had rooted, leaching data were obtained for seven N applications scheduled across a 12-mo period.
Nitrate and Ammonium Leaching
The initial N application (23 Oct.) was applied based on the determination that all the species had sufficiently rooted. Nitrate concentration in the leachate peaked at 35 to 65 mg N L-1 8 d after application, but returned to very low levels (
0) by Day 25 (Fig. 1
, Table 2). There were significant differences among species for total loss of NO3-N (Table 3), ranging from a low of 24% of the applied N for St. Augustinegrass to a high of 56% for Meyer zoysiagrass. Hybrid and common bermudagrass also showed substantial leaching, averaging 52 and 47% loss, respectively. It should be noted that these values, expressed as percentage of total applied N, are equivalent to 48, 112, 104, and 94%, respectively, of the N applied as NO3, since NH4NO3 was the fertilizer source.

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Fig. 1. Nitrate-N concentration in the leachate from six warm season turfgrasses following the first application (23 Oct.) of ammonium nitrate. Values are means of four samples.
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Table 2. Peak and volume-weighted average NO3-N concentrations in the leachate from six warm-season turfgrass species following seven N applications.
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Ammonium enters into cation exchange reactions and is considered to be fairly immobile and unlikely to leach in most soils. However, there was considerable NH4 leaching from most of the species following the first N application, probably due to the low cation exchange capacity of the sand. Ammonium concentrations in the leachate (Fig. 2)
, which also peaked on Day 7, ranged from
5 mg N L-1 for St. Augustinegrass to
20 mg N L-1 for Meyer zoysiagrass and the two bermudagrass species. Total N loss as NH4-N ranged from
4% of the applied N for St. Augustinegrass to 16% for Meyer zoysiagrass (Table 3).

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Fig. 2. Ammonium-N concentration in the leachate from six warm season turfgrasses following the first application (23 Oct.) of ammonium nitrate. Values are means of four samples.
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The second N treatment was applied on 22 November, 30 d after the first fertilizer application. Peak NO3 concentrations occurred 20 to 30 d after application and ranged from 5 to 35 mg N L-1 (Fig. 3)
, with St. Augustinegrass and centipedegrass being the lowest, and Meyer zoysiagrass and the two bermudagrasses being the highest. For all species, total loss of NO3-N was greatly reduced compared with the first application (Table 3). For example, leaching loss from centipedegrass and St. Augustinegrass following the second N application was reduced 86 and 92%, respectively, compared with the loss following the first application.

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Fig. 3. Nitrate-N concentration in the leachate from six warm season turfgrasses following the second application (22 Nov.) of ammonium nitrate. Values are means of four samples.
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Ammonium was almost undetectable in the leachate following the second N application, with concentrations typically <0.5 mg N L-1. Total loss amounted to <0.2% of the applied N (Table 3). This is in striking contrast to the NH4 lost following the first N application, and suggests either that nitrifying bacteria populations had significantly increased, or that the young root systems had developed to a degree where they were very efficient at absorbing the applied NH4. Several studies have shown that turfgrass roots absorb NH4 rapidly compared with NO3 (Bowman et al., 1989, 1998).
Several factors may explain the differences in N loss noted between the first and second N applications. First, the grasses had likely developed a deeper and more extensive root system by the second application, which would allow for more efficient N absorption. Bowman et al. (1998) measured less NO3 leaching from a deep-rooted compared with a shallow-rooted bentgrass. Similarly, Brauen et al. (1994) reported diminished NO3 leaching with increased rooting in creeping bentgrass. Second, a larger microbial biomass may have been present, which would contribute to greater N immobilization. Bigelow (1999) observed that microbial populations develop very quickly in new sand-based putting greens. Third, there was less water percolating through the columns following the second application which would cause the applied N to remain in contact with the root system for a longer period. The lower leaching fraction (Table 4) and concentration peak at 20 to 30 d after N application (Fig. 2) compared with 8 d for the first application is evidence of this.
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Table 4. Leaching fraction from six warm-season grasses. Data are averages for the interval between consecutive N applications.
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Following the third N application (21 March), NO3 concentrations and cumulative N loss were the lowest of any N application (Fig. 4 ; Tables 2, 3). Peak NO3 concentrations ranged from 0.2 to 5.7 mg N L-1, while volume weighted averages ranged from <0.1 to 2.0 mg N L-1. St. Augustinegrass was the most efficient and Meyer zoysiagrass the least efficient species at reducing N leaching, and this ranking remained consistent throughout the remainder of the study. Cumulative N loss was also reduced compared with the first two N applications, with losses ranging from 0.04 to 6.9% of the applied N (Table 3). These data reflect increased N efficiency by the turfgrasses, probably due to more extensive root growth or more rapid N uptake reflecting higher growth demand for N. Day length and light intensity were increasing during this period, and growth responded in parallel.

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Fig. 4. Nitrate-N concentration in the leachate from six warm season turfgrasses following the third application (21 Mar.) of ammonium nitrate. Values are means of four samples.
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Results from the final four N applications were remarkably consistent and illustrate genotypic differences in NO3 leaching potential (Tables 2, 3). St. Augustinegrass was clearly the best and Meyer zoysiagrass the worst species for controlling NO3 leaching. While peak NO3 concentrations and total N losses were fairly high, the data must be interpreted in the context of the substantial leaching fractions (3980%) maintained throughout this study. Several studies have identified irrigation practices, and specifically the amount of water moving through the rootzone, as a primary determinant of N leaching (Snyder et al., 1984; Morton et al., 1988; Bowman et al., 1998). It thus seems likely that leaching losses in this study would approximate a worst-case situation, and are not representative of losses expected under field conditions.
Plant Growth
Cumulative clipping production for each species was calculated for the 4 mo of April through July, 1997, corresponding to the period of most rapid growth by the six grasses. St. Augustinegrass produced significantly more leaf growth than the other species (Table 5), while the two bermudagrasses produced the least. There were also significant differences in N concentration in the clippings averaged during the same period, with the bermudagrasses having the highest and centipedegrass the lowest levels. Tissue N was negatively correlated with total clipping production (r = -0.92, P = 0.009), which seems somewhat counterintuitive since higher N is normally associated with greater growth. This relationship may indicate that warm season grasses differ fundamentally in the cellular N concentration required for productive function, with centipedegrass having the highest and the bermudagrasses the lowest N use efficiency.
Shoots and roots were harvested at the end of the 12-mo study to compare root architecture and biomass allocation among the species. Shoots comprised the majority of the biomass in all species (
83%, Table 5). Emerald zoysiagrass had the highest and common bermudagrass the lowest shoot mass. St. Augustinegrass had nearly twice the root mass of any other species, potentially increasing its ability to absorb NO3 from the soil system.
Nitrogen Recovery
Nitrogen recovery was estimated by traditional 15N labeling and by a longer-term analysis of N allocation to leaf tissue (Bowman et al., 2000). This latter method (Fig. 5)
integrates N acquisition, allocation, and remobilization across an extended period, and assumes that root and verdure biomass are constant across time. Cumulative N harvested in the clippings is plotted against time in days, and the slope of the linear regression represents average daily N allocation to leaf production. Comparing the slope to the average amount of N applied (also expressed on a daily basis) provides an estimate of N recovery during the longer term. This method is ideally suited for turfgrass systems, since it depends on frequent harvests. However, it may underestimate N recovery if N is partitioned to new verdure or root biomass.

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Fig. 5. Example of cumulative N harvest data analysis for estimating long term N use efficiency. Data are presented for centipedegrass and St. Augustinegrass, representing the least and most efficient species, respectively.
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On the basis of this long-term analysis for the period of 22 May through 26 Aug., St. Augustinegrass had the most efficient recovery, allocating 84% of the applied N to new leaf tissue (Table 6). Centipedegrass was the least efficient, allocating only 63% of that applied.
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Table 6. Nitrogen absorption, based on recovery of applied 15N in plant tissues (short term) and long term N allocation to leaf growth, based on regression analysis of cumulative N harvested across time.
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Short-term N use efficiency, based on total recovery of 15N in all plant tissues, ranged form 63 to 84% of the applied N, essentially identical to the range of values from the long-term analysis (Table 6). On the basis of 15N analysis, hybrid bermudagrass was the most efficient and Meyer zoysiagrass the least efficient species for N absorption. The majority of labeled N was in the shoots (3863%) followed by clippings (539%) and then roots (36%). Both bermudagrasses had significantly greater amounts of N allocated to new leaves compared with the other species. The high percentage of N allocated to shoots compared with leaves suggests a storage function, and could be related to changes associated with the declining photoperiod in September and the onset of dormancy. Relatively little 15N remained in the roots of any species, which reflects the comparatively low biomass in the root systems of these turfgrasses.
These 15N data indicate that the warm season turfgrasses vary with regards to resource allocation. Centipedegrass and Emerald zoysiagrass, for example, stored most recently absorbed N in the shoots (60 and 63%, respectively) and presumably would partition it to new leaf growth across extended periods. By contrast, the bermudagrasses were somewhat more profligate and allocated a large portion of recently absorbed N to new leaves during a short period. This is supported by the temporal patterns in leaf tissue N (Fig. 6)
. Bermudagrass leaf N concentrations fluctuated dramatically in response to N application, in contrast with the relatively stable N concentrations in the other four species. This implies that bermudagrass is more likely to exhibit periodic growth cycles timed to N applications than are the other species. It also suggests that the bermudagrasses experienced periods of greater N deficiency relative to the other species, which could explain the low clipping yields noted above for these normally prolific species.

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Fig. 6. Concentration of reduced N in the clippings from six warm season turfgrasses during the final 6 mo of the study. Arrows indicate NH4NO3 applications (50 kg N ha-1). Values are means of four samples.
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Root Length Density
Root length density was measured at the end of the experiment as a function of soil depth. There were no significant differences in RLD among species at the 5- and 18-cm depths (Table 7). At soil depths >30 cm, St. Augustinegrass and the bermudagrasses had significantly higher RLD than the other species. Nitrate leaching loss, averaged across the final three N applications, was negatively correlated with RLD averaged across the 30- to 55-cm depth (r = -0.80, P = 0.04; Fig. 7)
. This is consistent with data documenting turfgrass root architecture as a determinant of NO3 leaching (Bowman et al., 1998) and uptake (Sullivan et al., 2000). Increased rooting depth and density may reduce N leaching in several ways. First, a pulse of applied N will be exposed to a deeper root system for a longer period of time as the N moves through the soil profile, allowing for more uptake. Second, a denser root system may support larger populations of soil microorganisms, which will contribute to N immobilization. Third, increased root density reduces the average distance between roots, which in turn reduces diffusional limitations to ion uptake. Increased N uptake limits the amount of fertilizer subject to leaching. And fourth, more extensive rooting contributes to drought avoidance and overall health and activity of the turf.

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Fig. 7. Nitrate leaching loss as a function of root length density. Leaching losses are averaged across the final three N application periods, and root length densities are averaged across the 30- to 55-cm soil depth.
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This research has documented genotypic differences among the warm-season turfgrasses for NO3 leaching potential, N absorption, and root architecture. St. Augustinegrass and both common and hybrid bermudagrass were the most effective at reducing leaching, while Meyer and Emerald zoysiagrass were significantly less effective. Nitrate and ammonium readily leached from all species shortly after planting, but leaching declined substantially as the root systems developed. Volume weighted NO3 concentrations in leachate from established turf were all below the 10 mg N L-1 threshold with a single exception of Meyer zoysiagrass following the June N application. These data must be considered in the context of experimental conditions chosen to maximize leaching potential and magnify differences among the species. Actual management and field conditions are unlikely to approach the worst case conditions used in this study, and leaching is thus likely to be considerably reduced from levels reported above. From a practical standpoint, the results indicate that species selection and rooting depth are primary determinants of NO3 leaching and N use efficiency, and highlight the importance of carefully managing both fertility and irrigation when establishing warm-season grasses from sod.
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NOTES
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Research supported by the North Carolina Agric. Exp. Stn. and the Turfgrass Council of North Carolina.
Received for publication November 27, 2000.
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M. J. Fagerness, D. C. Bowman, F. H. Yelverton, and T. W. Rufty Jr.
Nitrogen Use in Tifway Bermudagrass, as Affected by Trinexapac-Ethyl
Crop Sci.,
March 1, 2004;
44(2):
595 - 599.
[Abstract]
[Full Text]
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D. J. Lee, D. C. Bowman, D. K. Cassel, C. H. Peacock, and T. W. Rufty Jr.
Soil Inorganic Nitrogen under Fertilized Bermudagrass Turf
Crop Sci.,
January 1, 2003;
43(1):
247 - 257.
[Abstract]
[Full Text]
[PDF]
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