Published online 1 September 2007
Published in Crop Sci 47:2198-2209 (2007)
© 2007 Crop Science Society of America
677 S. Segoe Rd., Madison, WI 53711 USA
FORAGE & GRAZINGLANDS
The Long-Term Impact of Phosphorus and Potassium Fertilization on Alfalfa Yield and Yield Components
W. K. Berg,
S. M. Cunningham,
S. M. Brouder,
B. C. Joern,
K. D. Johnson,
J. B. Santini and
J. J. Volenec*
Dep. of Agronomy, Purdue Univ., 915 West State St., West Lafayette, IN 47907-2054
* Corresponding author (jvolenec{at}purdue.edu).
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ABSTRACT
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Addition of P and K fertilizer can increase alfalfa (Medicago sativa L.) yield and stand persistence, but the yield components associated with P- and K-induced variation in agronomic performance are not clear. Our objectives were: (i) to determine the impact of P and K nutrition on productivity of a relatively old alfalfa stand; and (ii) determine which yield components are associated with changes in alfalfa forage yield. Treatments were a factorial combination of four P and five K rates replicated four times. Forage harvests occurred four times annually. Plant populations were determined in early December and late May each year. When compared to unfertilized plots, addition of P and K increased forage yield each year. Fertilization with P decreased plants m–2 at all K application rates, but especially in plots fertilized with P, but not K. By comparison, plots fertilized with K, but not fertilized with P, had the higher plant population densities. Although regression analysis eventually revealed a positive association between forage yield and shoots m–2 in 2003 and 2004, the greatest forage yields were not obtained in plots with the greatest plant population densities, shoots plant–1 or shoots m–2. Regression and path analysis revealed that improved forage yield in P- and K-fertilized plots was consistently associated with greater mass shoot–1.
Abbreviations: ANOVA, analysis of variance
The Long-Term Impact of Phosphorus and Potassium Fertilization on Alfalfa Yield and Yield Components
W. K. Berg,
S. M. Cunningham,
S. M. Brouder,
B. C. Joern,
K. D. Johnson,
J. B. Santini and
J. J. Volenec*
Dep. of Agronomy, Purdue Univ., 915 West State St., West Lafayette, IN 47907-2054
* Corresponding author (jvolenec{at}purdue.edu).
Addition of P and K fertilizer can increase alfalfa (Medicago sativa L.) yield and stand persistence, but the yield components associated with P- and K-induced variation in agronomic performance are not clear. Our objectives were: (i) to determine the impact of P and K nutrition on productivity of a relatively old alfalfa stand; and (ii) determine which yield components are associated with changes in alfalfa forage yield. Treatments were a factorial combination of four P and five K rates replicated four times. Forage harvests occurred four times annually. Plant populations were determined in early December and late May each year. When compared to unfertilized plots, addition of P and K increased forage yield each year. Fertilization with P decreased plants m–2 at all K application rates, but especially in plots fertilized with P, but not K. By comparison, plots fertilized with K, but not fertilized with P, had the higher plant population densities. Although regression analysis eventually revealed a positive association between forage yield and shoots m–2 in 2003 and 2004, the greatest forage yields were not obtained in plots with the greatest plant population densities, shoots plant–1 or shoots m–2. Regression and path analysis revealed that improved forage yield in P- and K-fertilized plots was consistently associated with greater mass shoot–1.
Abbreviations: ANOVA, analysis of variance
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INTRODUCTION
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THE IMPACT of P and K fertilization on alfalfa yield and persistence has been studied for many years (Nelson and MacGregor, 1957; Markus and Battle, 1965; Collins and Duke, 1981). A comprehensive understanding of the physiological and morphological basis for changes in alfalfa productivity in response to P and K fertilization remains elusive, however, because the components of yield are rarely analyzed, instead studies have tended to focus on cultivar x nutrient interactions (James et al., 1995). Other single-nutrient studies were conducted in controlled conditions that minimize many abiotic stresses (Li et al., 1997; 1998), or occurred in the field over a relatively short two or three year time periods (Collins and Duke, 1981; Collins et al., 1986). Recent studies have shifted focus to other questions such as P placement, and have shown occasional advantages to band application of P when compared to broadcast applications (Teutsch et al., 2000; Koenig et al., 2003). While these studies have been valuable in understanding several aspects of alfalfa P and K management, our knowledge of how alfalfa responds to P and K application remains rudimentary. This paper extends our earlier findings (Berg et al., 2005) on the impact of P and K fertilization on forage yield and yield component analysis during Years 1 to 3 (1998 to 2000) to learn how productivity and persistence change as an alfalfa stand ages between Years 4 to 7 (2001 to 2004).
Alfalfa plant populations are greatest immediately following establishment, and decline with time (Volenec, 1999). Adequate K nutrition is thought to enhance stand longevity (Wang et al., 1953; Smith, 1975). For example, Hanson and MacGregor (1966) reported that alfalfa stands receiving K fertilizer after first cutting, and both K and P fertilizer in autumn, maintained a denser stand after 10 years when compared to plots fertilized with only P. Surprisingly, there was no clear association between yield and stand density; however, plant counts were not reported in that study so the actual impact of P and K on alfalfa population densities is not known. However, Collins et al. (1986) counted crowns and reported that stand density in K-fertilized plots declined 50% between Years 1 and 2, whereas a 73% decline in plant density occurred in unfertilized control plots.
While fertilization with P often increases forage yield, conflicting results exist regarding the effects of P on alfalfa stand density. Jung and Smith (1959) reported that P fertilization was essential for plant survival. In contrast, Sanderson and Jones (1993) reported that P fertilization reduced alfalfa plant population densities. During their two year study, alfalfa populations declined from 495 to 98 plants m–2 in plots provided 59 kg P ha–1 annually, whereas unfertilized stands contained 134 plants m–2 at the conclusion of the study. Collins et al. (1986) also reported decreased alfalfa populations in plots fertilized with P when compared to unfertilized control plots. Hanson and MacGregor (1966) reported that broadcast applications of K with P maintained more dense alfalfa stands than fertilization with P alone. Therefore, because P application may decrease alfalfa plant populations, the greater yield of P-fertilized plots should be reflected in increases in the other yield components of alfalfa (shoots plant–1; mass shoot–1).
Alfalfa yield has been positively associated with shoots area–1. This yield component recently has been suggested as a criterion to determine whether or not to keep an existing alfalfa stand. Undersander et al. (1998) indicated that stem densities above 592 shoots m–2 (55 shoots ft–2) do not limit forage yield, however, some yield reduction is expected at shoot densities between 431 (40 shoots ft–2) and 592 shoots m–2. They recommend replacing the stand when shoot densities decline below 431 shoots m–2. Using shoots plant–1 and plant m–2 data of Suzuki (1991), we estimated shoots m–2 and observed the highest forage yields occurred at the lowest shoot densities (212 shoots m–2).
Increased mass shoot–1 has consistently been associated with improved agronomic performance of alfalfa irrespective of whether greater yields resulted from genetic selection (Volenec, 1985), improved soil fertility (Li et al., 1997, 1998), or better insect control (Kitchen et al., 1990). Cooper et al. (1967) attributed increased mass shoot–1 with improved K fertility to an increased number of leaves per shoot. Others have associated high mass shoot–1 with rapid shoot elongation rate after harvest (Volenec, 1985). During the initial three production years of the study reported here, mass shoot–1 was consistently associated with increased forage yield (Berg et al., 2005). Plant population densities and shoots per area were unaffected by P and K fertility, or declined slightly even though yield was increased with P and K fertilizer application. However, interplant competition may have prevented fertility effects on shoot and plant densities from being detected during Years 1 to 3 (1998 to 2000). It is not known how alfalfa yield components interact to maintain high forage yield as plant populations naturally decrease with stand age, and how P and K fertilizer applications alter yield and yield components under these conditions.
Our hypothesis is that P and K fertilization is important to maintain high alfalfa yields, and because stands decline with age, yield will be influenced by different components than we found previously in a relatively young alfalfa stand (Berg et al., 2005). Our objectives were: (i) to determine the impact of P and K nutrition on productivity of a relatively old alfalfa stand, and (ii) determine which yield components are associated with changes in alfalfa forage yield.
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MATERIALS AND METHODS
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Field Design and Sampling
In April 1997, a 1.4-ha site at the Throckmorton Purdue-Agricultural Center located 15 km south of West Lafayette, IN (40°N and 87°W) was seeded to Pioneer Brand 5454 alfalfa. This site was selected for study because soil tests indicated low concentrations of extractable P (9 to 15 mg kg–1 Bray P1) and low to moderate levels of exchangeable K (108 to 138 mg kg–1 exchangeable K) (Vitosh et al., 1996; Berg et al., 2005). Fertility treatments (0, 25, 50, and 75 kg P ha–1; and 0, 100, 200, 300, and 400 kg K ha–1) were applied to 5- x 10-m plots in a factorial design replicated four times. Two replications (field blocks) were placed on a Drummer silty clay loam (fine-silty, mixed superactive, mesic Typic Endoaquoll), while the other two replicates (field blocks) were placed on a Lauramie silt loam (fine-loamy, mixed, active, mesic, Mollic Hapludalf). Phosphorus (as triple super phosphate) and K (as potassium chloride) fertilizers were applied in split applications; with one half the annual amount broadcast after the first forage harvest in late May, and the remainder broadcast after the last forage harvest in mid-September. Chemical control of potato leafhoppers (Empoasca fabae Harris) and alfalfa weevil (Hypera postica Gyll.) was done as necessary using cyfluthrin [cyano(4-fluoro-3-phenoxyphenyl)methyl 3-(2,2-dichloroethenyl)-2,2-dimethylcyclopropanecarboxylate)]. Plots were sprayed twice with sethoxydim [2-(1-ethoxyiminobutyl)-5-[2-(ethylthio)propyl]-3-hydroxycyclohex-2-enone] to reduce invasion of grasses. Monthly averages for maximum and minimum temperatures (air temperatures at 1.4 m above the soil surface; and bare soil temperatures at 0.1-m depth below the soil surface) and precipitation from January 2001 through December 2004 were obtained at the Agronomy Center for Research and Education located 15 km north of the experimental site (Table 1
).
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Table 1. Maximum (max.) and minimum (min.) air and soil temperatures and total precipitation (precip.) by month for each year of the study. Temperature data are averages of daily maximums and minimums observed for each month.
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Four forage harvests occurred each year at approximately 30-day intervals beginning in late May and concluding in early September as described previously (Berg et al., 2005). A flail-type forage chopper (Carter Harvester, Brookston, IN) was used to harvest a 1-m-wide swath from the center of each plot. A subsample of chopped forage was weighed, dried at 65°C for 48 h, re-weighed, and percent dry matter of the forage determined. Dry matter yield of each plot was calculated based on the forage fresh weight, area harvested, and percent dry matter of the forage. A subsample of 50 shoots was hand-collected at approximately six randomly selected sites within each plot before harvest, dried at 65°C, weighed, and mass shoot–1 determined. Shoots m–2 was calculated by dividing yield m–2 by mass shoot–1. Taproots in a 0.5-m2 area of each plot were excavated to a depth of 20 cm and counted in May and December of each year to determine plant population density (plants m–2). Excavated areas were positioned so that at least a 2-m-wide strip in the center of the plot remained intact for forage harvest. Shoots plant–1 was calculated for each May harvest by dividing shoots m–2 by plants m–2.
Statistical Analysis
Analysis of variance was performed on the yield components, and the P, K, and P x K treatment effects were partitioned into orthogonal polynomial contrasts as previously described (Berg et al., 2005). In this analysis the ORPOL function in PROC IML of SAS was used to obtain orthogonal polynomial coefficients. These values were assigned to dummy variables representing linear, quadratic, cubic, and quartic polynomial contrasts and their interactions. The REG procedure was run with these dummy variables as independent variables; where the Type II sums of squares are the sums of squares for the corresponding contrasts. Regression models incorporating terms for all significant polynomial contrasts were developed and evaluated on treatment means. Occasionally a regression model exhibiting an anomalous relationship was rejected. Differences in equation slopes were determined using the standard error of the difference between the slopes adjusted for the experimental error. Because forage yield is a linear function of mass shoot–1 and shoots m–2 (and at May harvests mass shoot–1, shoots plant–1, and plants m–2), and given that yield components themselves are often correlated with one another, we used path analysis (also known as analysis of correlation, Li, 1956) to determine the direct effects of individual yield components on forage yield. The ratio of path coefficients of the yield components at a particular harvest provides an estimate of the relative influence of each yield component on forage yield at that harvest. Statistical analyses were performed using SAS (SAS, 1999).
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RESULTS AND DISCUSSION
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Forage Yield
When compared to the unfertilized control plots, application of P and K resulted in greater forage yield each year of this study (Fig. 1
). Maximum forage yields were lower in 2003 when compared to the other years, possibly due to the lower soil temperatures in June and July of that year (Table 1). Evenson (1979) showed a linear reduction in herbage growth as crown temperatures declined below 30°C. Between 2001 (fourth year of production) and 2004 (seventh year of production), the impact of adequate P and K fertilization on forage yield became progressively more evident. In 2001 there was a 33% difference between the highest and lowest forage yield, but by 2004 this difference had increased to 300% (Fig. 1). These large changes in yield provide an excellent opportunity to quantify the impact of P and K fertility on yield and yield components of alfalfa.

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Figure 1. Influence of P and K fertilizer application on annual forage yield (kg ha–1 yr–1) in 2001 to 2004. Contour plots are generated from the regression equations listed in Table 3 for total yield.
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Table 3. Regression equations for estimating the influence of P and K on yield and yield components of alfalfa at Harvests (H) 1 to 4 in each year, 2001 to 2004.
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Significant orthogonal polynomial contrasts existed for K1 and K2 for yield at each harvest (Table 2
). Significant contrasts also were found for P1 and P2 for total yield and Harvest 1 of each year, and all other harvests except Harvest 4 in 2001; Harvest 3 and 4 in 2002; Harvest 4 in 2003; and Harvests 2 and 3 in 2004. The polynomial orthogonal contrast for the K1 x P1 interaction was significant at most harvests in 2001 and 2002, and every harvest of 2003 and 2004. This interaction indicates that application of both P and K fertilizers generally resulted in higher forage yield than when either nutrient was added alone (Fig. 1). This agrees with the initial three production years in this trial (1998 to 2000) where highest forage yields were obtained with application of both P and K fertilizer rather than a single nutrient applied alone (Berg et al., 2005). Hanson and MacGregor (1966) also reported that repeated application of both P and K were necessary to obtain high alfalfa yields during the 10 year life of a stand.
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Table 2. Level of significance for the K and P treatment effects and significant orthogonal polynomial contrasts on individual harvest and annual yield in 2001, 2002, 2003, and 2004.
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As expected, the lowest forage yields in 2001 were obtained in the unfertilized control plots. In 2002 and 2003 both unfertilized control plots and those fertilized with 75 kg P ha–1 yr–1 without K fertilization had the lowest forage yields. Surprisingly, in 2004 yield of plots fertilized with 75 kg P ha–1 yr–1 without K fertilization yielded less than the unfertilized control plots. During the 10 year life of a stand, Hanson and MacGregor (1966) reported that plots left unfertilized and those fertilized with only P or N, but not K, had the lowest forage yield. James et al. (1995) reported that annual applications of K fertilizer were necessary to maintain high forage yield, whereas application of 110 or 220 kg P ha–1 was sufficient to maintain high yield for at least three years.
Yield Components
Plant Population Density
Alfalfa forage yield can be described as the product of three components: plants area–1, shoots plant–1, and mass shoot–1 (Volenec et al., 1987). Ignoring the influence of P and K, plots averaged 130 plants m–2 in May 2001, and this decreased to 46.7 plants m–2 by May 2004 (Table 3
; intercepts of regressions). Analysis of variance indicated significant linear effects of K on plants m–2 in five of eight samplings (Table 4
). Where K effects were significant, plants m–2 increased with K fertilization (Table 3). Analysis of variance revealed significant linear and quadratic effects of P on plants m–2 at six of eight samplings between May 2001 and December 2004 (Table 4). Where significant, application of P reduced plants m–2 (Table 3). At the highest rates of P (75 kg ha–1) and K (400 kg ha–1) the regressions in Table 3 predicted an increase of 17 plants m–2 due to K fertilization, and a reduction of 27 plants m–2 due to P averaged across the five of eight harvests where both P and K significantly influenced plants m–2.
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Table 4. Levels of significance for the K and P treatment effects and orthogonal polynomial contrasts on individual plant population densities (plants m–2).
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Plant death was seasonal with few losses occurring over winter (December to May). Virtually all reductions in plants m–2 occurred between the May and December taproot samplings. Averaged over fertility treatments, plant population densities declined 27, 27, and 24 plants m–2 between May and December of 2001, 2002, and 2003, respectively, while 6, 0, and 0 plants m–2 died over the winters of 2001 to 2002, 2002 to 2003, and 2003 to 2004, respectively. It is possible that winter-injured plants survived through Harvest 1, taken in May of each year, but subsequently died later in summer.
Adequate K nutrition has often been positively associated with improved alfalfa persistence (Gerwig and Ahlgren, 1958; Burmester et al., 1991; Simons et al., 1995). However, K fertilizer application enhanced plant populations in only five of eight samplings, where on average, the 400 kg K ha–1 yr–1 treatment increased populations by 28% (Table 3; calculated using regression equations). By comparison, P fertilization reduced plant populations in six of eight samplings by an average of 48% at the 75 kg P ha–1 fertilizer rate (Table 3). The decrease in alfalfa populations observed in response to enhanced P fertilization agrees with trends previously reported (Sanderson and Jones, 1993; Berg et al., 2005). In a decade-long alfalfa fertility study, Hanson and MacGregor (1966) reported that stand densities in plots fertilized with both P and K were greater than stand densities of plots fertilized with P alone. In addition, they reported no association between variation in stand density (37 to 62% of stand) and forage yield. We found that the highest rates of P and K fertilization resulted in the greatest forage yields in 2004 (Fig. 1), and these occurred at intermediate plant populations of approximately 40 plants m–2 (Fig. 2
). Although 50% higher plant populations occurred when plots were fertilized with K but not P (Fig. 2), the relatively small size of these P-limited plants resulted in lower forage yield (Fig. 1).

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Figure 2. Influence of P and K fertilizer application on shoots plant–1, shoots m–2, plants m–2, and mass shoot–1 at Harvest 1 in May 2004. The contour plots were generated from the regression equations listed in Table 3 for Harvest 1 of 2004.
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Plant population is often inversely related to plant size in natural communities. Using a broad range of species, Yoda et al. (1963) consistently found a –3/2 slope (–1.5) when log(plant mass) is plotted against log(plant density). Harvest 1 data from each year were analyzed to determine the influence of P and K nutrition on the plant size- density relationships in alfalfa. These data were selected for analysis because the May plant counts enabled us to calculate individual mass plant–1. For plots provided 0 kg K ha–1 yr–1 the slope of the regression of log(plant mass) versus log(plant density) ranged from –0.87 to –0.99 (Table 5
). This indicates that decreases in plant population were not offset by the expected increases in plant size (i.e., slopes were not –1.5); a possible result of K-limited growth. By comparison, the slope of the log(plant mass) versus log(plant density) regression for plants provided 400 kg K ha–1 yr–1 had slopes that averaged –1.25 over the four years of the study. This indicates that compensatory increases in plant size occurred as a result of reductions in plant population if high K fertilization rates were provided. The log(plant mass) versus log(plant density) slopes for the other K-fertility treatments were generally intermediate in their responses (Table 5). In the first three production years of this study (1998 to 2000) slopes of the log(plant size) versus log(plant density) relationships ranged from –1.3 to –1.9, with the 0 kg K ha–1 yr–1 plots having a slope of –1.5 (Berg et al., 2005). At that time soil test K levels, though low, may have been sufficient to permit alfalfa growth to occur as expected in response to reductions in plant populations.
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Table 5. Plant populations, regression equation slopes of the log above ground plant weight as a function of log plant population, and plant mass at the May Harvest (Harvest 1) in 2001, 2002, 2003, and 2004. Potassium treatment data are averaged over the P treatments, while data tabulated for the P treatments are averaged over the K treatments.
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The slopes of the log(plant size) versus log(plant density) regressions for the P-treatments changed with time and P rate (Table 5). Slopes ranged from –0.82 to –0.93 in 2001 (except 50 kg P ha–1 yr–1) and increased to about –1.3 in 2002 (except –1.1 for the 25 kg P ha–1 yr–1). Thereafter slopes of the P-fertilized plots became less negative, especially for plots fertilized with 50 and 75 kg P ha–1 yr–1. This pattern of less negative slopes with increasing rate of P fertilizer also was observed in 1998 to 2000 of this study (Berg et al., 2005), and indicates that at a given population plants provided 50 and 75 kg P ha–1 yr–1 have greater than anticipated plant mass (Table 5).
These results agree with those of Lemaire et al. (1991) who suggested that competition for light within an alfalfa stand decreased plant populations in a manner consistent with the –3/2 thinning law. Preferential death of the smallest plants in the population occurred because of extensive shading and low plant N content. Kays and Harper (1974) interpreted changes in slope of the log(plant size) versus log(plant density) regression to values greater than –1.5 (i.e., –1.0) as an indication that the stress factor (in our case, K and P deficiency) has a causal role in stand thinning in addition to interplant competition per se. This highlights the importance of adequate K and P nutrition in enabling increases in plant size to offset reductions in plant population that are essential for maintaining high forage yield as alfalfa stand density declines.
Shoots Plant–1
Taproots were excavated and counted only at Harvest 1, therefore, shoots plant–1 can only be calculated at this harvest of each year. Shoots plant–1 was not influenced by P or K fertilization in 2001, 2002, or 2003; however, in 2004 significant orthogonal polynomial contrasts were obtained. The ANOVA (analysis of variance) of shoots plant–1 in 2004 revealed that while P fertilization increased shoots plant–1, K fertilization reduced shoots plant–1 at most P rates (Fig. 2). For example, plants provided 75 kg ha–1 P had 14 shoots plant–1 when fertilized with 0 kg K ha–1 yr–1, but shoots plant–1 declined to between six and eight shoots plant–1 when plants were fertilized with 75 kg ha–1 P and 300 or 400 kg K ha–1 yr–1. The highest shoots plant–1 values (14 shoots plant–1) occurred in plots with the fewest plants m–2 (20 plants m–2) (Fig. 2). This agrees with previous results where genetic differences in shoots plant–1 were only evident when plant populations declined to 22 or fewer plants m–2 (Volenec et al., 1987).
While shoots are clearly needed for forage yield, high forage yield was not closely associated with greater shoots plant–1. For example, in 2004, the greatest yield was produced in plots provided 400 kg K ha–1 yr–1 and 75 kg P ha–1 yr–1 (Fig. 1) that had 7 or 8 shoots plant–1 (Fig. 2), whereas plots fertilized with 75 kg P ha–1 yr–1 and no K fertilizer had the highest number of shoots plant–1, but the lowest forage yield in 2004 because of high plant mortality. During the initial three production years of this study (1998 to 2000), slight increases in shoots plant–1 occurred as plant populations decreased, but these changes were independent of P and K fertilization (Berg et al., 2005). These findings also have implications for alfalfa improvement strategies because genetic enhancement of shoots plant–1 is unlikely to result in higher yielding alfalfa because genetic differences in shoots plant–1 are not evident under competitive conditions found in most stands (Volenec et al., 1987)
Shoots m–2
The ANOVA revealed an occasionally significant K x P orthogonal polynomial contrast (Table 6
). Potassium had a significant positive influence on shoots m–2 in 11 of 16 harvests, no significant impact on two of 16 harvests, and a significant negative effect in three of 16 harvests (Table 3). Using regression equations in Table 3, the positive response to 400 kg ha–1 yr–1 K fertilization rate ranged from a low of 19 shoots m–2 (Harvest 2 of 2001) to a high of 182 shoots m–2 (Harvest 3 of 2004), with an average positive response of 94 shoots m–2. At the 400 kg K ha–1 yr–1 rate the three negative responses of shoots m–2 to K fertilizer application ranged from a reduction of 90 shoots m–2 (Harvest 1 of 2001) to a reduction of 152 shoots m–2 (Harvest 4 of 2002), with an average reduction of 112 shoots m–2.
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Table 6. Level of significance for the K and P treatment effects and orthogonal polynomial contrasts on individual harvest for shoots m–2.
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By comparison, fertilization with P had no impact on shoots m–2 at seven of 16 harvests, enhanced shoots m–2 at one harvest (Harvest 3 of 2003: regression equation in Table 3 predicted 17 additional shoots m–2 at the 75 kg P ha–1 yr–1 rate), and reduced shoots m–2 at the other eight harvests including an average reduction of 68 shoots m–2 at Harvest 1 of each year (predicted using the regression equations in Table 3). Shoots m–2 was least responsive to P fertilization at Harvest 4, where there was no significant effect of P fertilization in three of four years.
Regression analysis of shoots m–2 versus forage yield revealed significant linear relationships between these two traits at 14 of 16 harvests (Table 7
). The r2-values ranged from 0.00 (Harvest 1 of 2002) to 0.94 (Harvest 2 of 2003). Although the r2-values were relatively low, the negative slopes obtained in three of four harvests in 2001 indicate that increases in shoots m–2 reduced forage yield at these harvests. Harvests 2 and 3 in 2002, and all harvests in 2003 and 2004 exhibited a positive relationship between shoots m–2 and forage yield indicating that increasing shoots m–2 at these harvests would have a positive impact on forage yield. The negative relationship between forage yield and shoots m–2 that was observed at three of four harvests in 2001 confirms a pattern also observed during 1998 to 2000 where fertility-driven increases in forage yield coincided with reductions in shoots m–2 (Berg et al., 2005). Although a positive relationship between shoots m–2 and yield also was evident at every harvest in 2003 and 2004, the greatest forage yields were never obtained in plots with the highest shoot densities. For example, in May 2004 shoot densities ranged from 200 shoots m–2 in plots provided 75 kg P ha–1 yr–1 and no K to 340 shoots m–2 in plots fertilized with 300 or 400 kg K ha–1 yr–1 without P (Fig. 2). The highest forage yield was obtained at shoot densities of approximately 260 shoots m–2 that occurred when high rates of both P and K were provided (Fig. 1). Despite this inconsistency, alfalfa yield potential is often predicted using shoots area–1 with significant losses in forage yield anticipated when shoot density declines below the "benchmark" value of 430 shoots m–2 (40 shoots ft–2) (Undersander et al., 1998). The high forage yields obtained in May of 2004 occurred at shoot densities far below this benchmark value (Fig. 2). Additional research is needed to accurately define critical shoot densities for high yielding alfalfa and how this value varies with P and K fertility, harvest, and stand age.
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Table 7. Regression equations, r2-values, and P-values for the influence of shoots m–2 and mass shoot–1 (g) on forage yield at each harvest in 2001, 2002, 2003, and 2004.
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Mass Shoot–1
Analysis of variance revealed significant linear and quadratic effects of P and K on mass shoot–1 (Table 8
). In addition, the K-linear x P-linear interaction was significant for mass shoot–1 at every Harvest in 2001 and 2004, and Harvests 1 and 4 of 2002 and 2003. Higher-order interactions were also occasionally significant. Using the regression models in Table 3 the predicted response of mass shoot–1 to application of 75 kg P ha–1 yr–1 ranged from –0.11 (Harvest 2 of 2004) to 0.15 g shoot–1 (Harvest 1 of 2001 and Harvest 2 of 2003), with a mean response over all years and harvests of 0.021 g shoot–1. This mean represents a 5% increase over the 0.445 g shoot–1 average predicted for the unfertilized control plots (mean of the y-intercepts from all year-harvest combinations in Table 3). Likewise for the 400 kg K ha–1 yr–1 treatment, the predicted responses ranged from a low of –0.16 g shoot–1 (Harvest 1 of 2003) to a high of 0.32 g shoot–1 (Harvest 2 of 2004), with a mean response over all year-harvest combinations of 0.088 g shoot–1. This mean represents a 20% increase over the average 0.445 g shoot–1 for the unfertilized control plots. The P x K interactions in the regression models resulted in consistently positive linear effects on mass shoot–1 ranging from a low of 0.12 g shoot–1 (Harvest 3 of 2002) to a high of 0.84 g shoot–1 (Harvest 1 of 2003), with a mean predicted response of 0.304 g shoot–1 (predicted using the regression equations in Table 3). This represents a 68% increase over the average 0.445 g shoot–1 for the unfertilized plots, and emphasizes the synergistic effects of providing adequate levels of both nutrients over high rates of either nutrient alone when the soil is deficient in both P and K.
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Table 8. Level of significance for the K and P treatment effects and orthogonal polynomial contrasts by individual harvest for mass shoot–1.
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The influence of P and K on mass shoot–1 was similar to the effects of P and K on total forage yield. Regression analysis of mass shoot–1 on forage yield revealed a significant, consistently positive relationship between these two traits at all harvests each year (Table 7). The r2-values ranged from 0.19 (Harvest 1 of 2004) to 0.74 (Harvest 4 of 2001 and Harvest 1 of 2002), and were greater than 0.45 in 11 of 16 harvests indicating that much of variation in forage yield at most harvests was associated with mass shoot–1. By comparison, the r2-values from regression of shoots m–2 on forage yield were greater than 0.45 less frequently, only six of 16 harvests (Table 7).
Mass shoot–1 was consistently positively associated with forage yield at every harvest in each year (Table 7). Like forage yield, mass shoot–1 consistently responded to P and K fertilization, including significant interactions for P and K at nearly every harvest each year (Table 8). Subsequent analysis of all 28 harvests obtained in this seven year study revealed no evidence for a decline in the response of yield to increased mass shoot–1 across the range in values (0.2–3.0 g shoot–1) (Volenec et al., 2005). Previous research demonstrated genetic variation in mass shoot–1 and showed that this trait was negatively correlated with genotypic differences in shoots plant–1 (Volenec et al., 1987). Additional research is needed to understand the genetic constraints on mass shoot–1 and determine if the negative correlation between shoots plant–1 and mass shoot–1 observed in this study will serve as an impediment to alfalfa yield improvement using yield component strategies.
Both mass shoot–1 and shoots m–2 were positively associated with forage yield at some harvests (Table 7). However, these two yield components are often negatively correlated to one another, which confounds the interpretation of correlation analysis. These two yield components also are mathematically related because we calculate shoot population densities based on forage yield and shoot mass measurements. To better understand the direct effect of each yield component on forage yield, two-way path analysis (shoots m–2 and mass shoot–1) was performed at all harvests, and three-way path analysis (plants m–2, shoots plant–1, and mass shoot–1) was performed using data from Harvest 1 of each year where we had plants m–2 information. Plant breeders have used path analysis to understand the relative importance of yield components of grain crops to grain yield. The common approach is to calculate the ratio of the path coefficients because this ratio estimates the relative contributions of individual yield components to yield at that harvest (Li, 1956; Dofing and Knight, 1992; Gravois and Helms, 1992).
Two-way path analysis revealed that mass shoot–1 generally made a greater contribution to forage yield in this study than did shoots m–2. At 13 of 16 forage harvests the ratio of path coefficients, mass shoot–1:shoots m–2, exceeded 1 indicating a greater relative direct effect of mass shoot–1 on forage yield (Table 9
). At Harvests 3 and 4 of 2004 the ratio of these path coefficients was approximately equal to 1 indicating that these two yield components were of equal relative importance in determining forage yield near the end of the study. Only at Harvest 2 of 2003 was shoots m–2 of greater importance than mass shoot–1 in determining forage yield of this alfalfa stand. At Harvest 1 of each year the ratio of path coefficients was 2.2 or greater indicating that at this harvest, mass shoot–1 was at least twice as important as shoots m–2 in directly affecting forage yield.
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Table 9. Path coefficient analysis at each harvest in every year (2001 to 2004). The direct effects of yield components on forage yield are shown along with the ratio of path coefficients so the relative impact of these components on yield can be estimated.
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Three-way path analysis (mass shoot–1; shoot plant–1; plants m–2) obtained at Harvest 1 of each year also revealed that mass shoot–1 had a greater direct effect on forage yield relative to the other yield components (Table 9). In this three-way analysis, mass shoot–1 had at least a 3.8-fold greater direct effect on forage yield than did shoots plant–1. In May 2002 and 2003, mass shoot–1 had fourfold greater impact on yield than either shoots plant–1 and plants m–2, which were similar in their relative impact on yield. In May 2004, the plants m–2 was twice as important as shoots plant–1 in determining forage yield (ratio of 4.1:0.5:1.0 for mass shoot–1:shoots plant–1:plants m–2, respectively) indicating that plant populations were at least as important as shoots plant–1 in determining forage yield in 2002 to 2004. Although path analysis was not routinely used to analyze alfalfa yield components, our findings agree with results of Frakes et al. (1961) who reported that stem and leaf weight (components of mass shoot–1) had much greater direct effects on alfalfa yield than did stem number (shoots plant–1). Thus, cultivars with the genetic potential to produce large shoots versus many shoots per crown would be expected to be more responsive to P and K fertilizer applications and have high yield under well-fertilized conditions.
Both regression and path analysis revealed that mass shoot–1 of alfalfa was consistently related to improved forage yield in response to P and K fertilization. This conclusion agrees with our observations from 1998 to 2000 (the first three production years of this stand) when mass shoot–1 also was consistently associated with forage yield (Berg et al., 2005). Factors that increase mass shoot–1 including genetics, environment, and management are likely to result in higher forage yield of alfalfa. From a plant development perspective, increased mass shoot–1 results from two plausible mechanisms: rapid initiation of new shoots from crowns after forage harvest; and high shoot elongation rates following initiation of growth (Singh and Winch, 1974; Li et al., 1997). We have previously shown that genetic variation in shoot elongation rate exists in alfalfa, and that those cultivars and germplasms with high shoot elongation rates had greater mass shoot–1 and high forage yields (Volenec, 1985).
We also observed that as stands aged increasing rates of P, and especially K were required to maintain high forage yield. Current work is focusing on how soil test P and K concentrations changed between 1998 and 2004, and how this influenced tissue P and K concentrations and the decline in agronomic performance of alfalfa over time. Additional studies are underway to understand the impact of long-term P and K fertilization on alfalfa forage quality (Lissbrant et al., 2006a). Preliminary results indicate that concentrations of neutral detergent fiber, acid detergent fiber, and lignin are slightly lower, and crude protein concentrations higher in forage of P- and K-deficient alfalfa plants. In addition, we are using cluster analysis to better understand how P and K fertilization influence alfalfa yield and persistence over time (Lissbrant et al., 2006b), and what role organic reserves in alfalfa taproots have in alfalfa yield and persistence to develop diagnostic tools to better predict long-tem alfalfa performance.
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NOTES
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All rights reserved. No part of this periodical may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage and retrieval system, without permission in writing from the publisher. Permission for printing and for reprinting the material contained herein has been obtained by the publisher.
Received for publication September 13, 2006.
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