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Foxtail millet shows significant genotypic variation for all the traits under field conditions

Analysis of variance revealed that significant genotypic variation existed for all the traits under both P- and P+ conditions (Table 1). The season and genotype x season component also showed non-significant variance. The heritability and error variance showed profound influence of genotypic variance on the total variance for all the traits in the population. These estimates are indicative of the general trends in the influence of different sources of variation on the heritable component of the trait [44].

Cumulative ranking of foxtail millet genotypes for all the significant traits under field conditions

Based on the cumulative ranking, genotypes ISe and ISe were in the top 5 in both Pi fertilized and unfertilized field plots (Fig 1A). Other genotypes that performed well under both P- and P+ conditions were CO7 (4^th, 7^th), ISe (10^th, 2^nd), ISe (7^th, 3^rd) and ISe (5^th, 14^th). Other top genotypes were ISe (3^rd under P-) and ISe and ISe (2^nd and 4^th under P+, respectively). Maxima followed by ISe and ISe showed poor performance in the field under both P- and P+ conditions. The other genotypes that were poor under P- are ISe and ISe

Rank sums of genotypes based on (A) field and (B) greenhouse evaluation. The 10 extreme genotypes are selected under each P regime, P+ and P-. P+ ranks are ordered from poor (having high-rank sum) to good (having low-rank sum). P ranks are ordered from good (low-rank sum) to poor (high-rank sum). Five genotypes with the lowest rank sum in each group are selected as best genotypes (green), and five genotypes each with highest rank sum are selected as poor genotypes (red).

Foxtail millet genotypes show differential responses to Pi under natural field and greenhouse conditions

The mean values of the responses of each trait for all the genotypes analysed in the field experiments are given in Table 2 and S2 Table. When the good and poor performing genotypes were compared, based on the rank sums for the P+ plot, statistically significant differences were seen for all the traits. Even in the case of P- conditions, conspicuous differences were recorded for all the traits. However, when the selection was done for extreme five genotypes in both the directions in both the treatments, differences in phenotypic performance were found to be reduced.

In total five traits were analyzed in greenhouse grown plants (Fig 1B). ISe , which was the highest-ranking genotype under field conditions in both Pi regimes, ranked low when grown under greenhouse conditions. Maxima, which showed a poor field performance under both fertilized and unfertilized conditions, was the fourth best performer under P+ treatments, and mid-ranking under P- in the greenhouse. Similarly, CO 6, which performed very well under both P+ and P- in the greenhouse experiments, indicated a moderate response under field conditions (37^th under P- conditions and 24^th under P+ conditions). ISe performed well under both field as well as greenhouse conditions and was the most consistent high-responding genotype in the study. Other high performers were CO7 and ISe Further, the genotype CO5 which was placed at the 13^th position in both P- and P+ soil, was placed at the first position under P+ and 6^th position under P- situations under greenhouse evaluation.

Greenhouse-grown plants also showed a high degree of variation under differing levels of Pi (S3 Table). Genotype ISe had the lowest rank sum (high performer) under P-, followed by CO6, ISe , ISe and ISe (Fig 1B). Genotypes ISe and ISe had the highest (low performer) rank sum under P-. Under high phosphate, the cumulative rank order was very different. The genotype CO6 was 2nd under P+. Genotype ISe performed poorly under P+ as well as P- (Fig 1B). Collectively, the high responding genotypes under P- conditions had significantly higher biomass, SL, RL, RHD and RHL than low performers (Table 3). The low performer, ISe , registered significantly higher RHD and RHL than other low performers under P- condition, whereas RHL and RHD are comparable to those of high performers.

Root hair density and root hair length are associated with better PUE

Under P+ conditions, the five low performers were comparable to the five low performers under P- conditions for all parameters. Genotypes ISe , ISe and ISe were common to both treatments as low performers. ISe , ISe and ISe , grown under P+ treatment as well as under P- condition, lacked root hairs (Table 3 and Fig 2). In contrast, the root hairs in the other genotypes (ISe , ISe , ISe and ISe ) shown in Fig 2 were sparse or indiscernible under P+ conditions but were increased in length and number in P-conditions. The high responding genotypes under P+ were different from the high responding genotypes under P- with the exception of the local genotype CO6, which was ranked 2^nd under both P- and P+ (Fig 1B). Under P- conditions, CO6 attained 90% of the SL and 76% of biomass seen under P+. For the root traits under P-, CO6 attained 69% of the RL, % of the RHD and % of the RHL attained under P+ conditions.

Root hair images of selected genotypes of foxtail millet showing a response to P- and P+ in the greenhouse.

The image was taken after 15 days of growth under P- and P+ in the greenhouse. The genotypes ISe and ISe are on the top in the cumulative ranking and high responding genotypes for root hair formation under P- condition. The genotypes ISe , ISe and ISe are low performers under P- in greenhouse. The genotype ISe is a low performer under P+ in greenhouse. The genotype ISe is an intermediate responder in both P- and P+.

All the high responding foxtail millet genotypes (ISe , CO6, ISe , ISe and ISe ) produced abundant and long root hairs under P- in the greenhouse experiment. RHL and RHD were associated with higher Pi content under P- conditions and Pi content is a good indicator to gain better information on plant response under low P conditions. We also assessed Pi uptake in selective genotypes which confirmed that RHL and RHD were associated with Pi uptake under P- conditions. Fig 2 shows the extent of variability of the root hairs in the high and low responders under P- conditions in greenhouse experiments.

P content of root and shoot tissues of high-responding genotypes were higher than those of low performers under field and greenhouse conditions

The total P and Pi contents were assayed in the leaf and root tissues of each of the five high and low responding genotypes under P- and P+ conditions, grown in the greenhouse (Fig 3) as well as in the field (Fig 4). As expected, in both high and low performers, more plant Pi is present under P+ conditions (Fig 3A and 3B) than under P- conditions (Fig 3C and 3D). Good responders under the P+ condition in the glasshouse generally had much higher ( fold higher) total leaf P and leaf Pi contents than low performers. Good responders on P- maintained similar levels of tissue P as poor responders on P+ suggesting higher PAE (compare Fig 3B and 3C). Under P-, the high responding genotypes maintained higher levels of both root and shoot P compared to poor responders (Fig 3C and 3D).