Above ground dry biomass and grain yield
Above ground biomass and grain yield were both significantly influenced by location. The above ground biomass was higher at Mokopane (3 694 kg/ha) followed by Dalmada (2 800 kg/ha) and the least biomass was obtained at Masemola with 1 557 kg/ha (Fig 2). A similar trend was also observed with grain yield where the yield at Masemola was the lowest at 363 kg/ha. These results showed that despite the availability of resources to the three emerging farmers, the productivity levels at the three farms differed significantly. The grain yield at Masemola was less than 30% of the yield achieved by the other two farmers. This could be attributed to a combination of factors, mainly the chilling damage observed during pod filling as well as poor management practices, particularly irrigation management.
When tropical crops are subjected to low temperatures, photosynthetic activities related enzymes are affected leading to low growth rate or even cessation of growth
(Charrier et al., 2015). The severity of the damage depends on the crop health status and the soil moisture (Niwas and Khichar, 2016). It was observed during data collection, even before the chilling damage, that the crop was experiencing some water stress. Weed management was also poor, probably due to lack of labour and the relatively larger area planted (4 ha) compared to the other farmers, compromising the crop nutrition status. The combination of a drying soil and the generally poor health status of the crop might therefore have contributed to extent of the chilling damage observed at Masemola. The generally poor crop management at Masemola was evidenced by the low biomass observed at flowering (Fig 2). Challenges observed at Masemola are similar to those reported for common bean farmers of Maharashtra in India
(Jaybhay et al., 2018). However, the grain yield observed in this study particularly at Dalmada and Mokopane were within range of yields observed in the country. South Africa dry bean yields are between 1.5 – 3.0 tons/ha. The yields are nevertheless obtained through nitrogen fertiliser applications of upto 50kgN/ha.
Effect of location and DAP on NDVI, SPAD value and LAI
To understand the variation in biomass and grain yield observed between the locations, some critical physiological parameters such as the SPAD value, NDVI, LAI and gas exchanges were determined. NDVI is a measure of the crop’s vigour while the SPAD value measures the greenness of the plant and is closely related to chlorophyll content. NDVI showed no differences between Dalmada and Mokopane even though Mokopane showed a tendency to have relatively higher values. The SPAD value showed significant variation between the locations at 45 and 75 DAP. At 45 DAP, SPAD value was higher at Masemola compared to the other locations but 30 days later the SPAD value at Masemola had the lowest value. On the other hand, the chlorophyll content, did not show any variation between the locations at 45 and 60 DAP but only at 75DAP. At 75 DAP, Mokopane had twice as high chlorophyll content compared to those observed at the other two locations. The findings of this study showed interesting results in that NDVI values at Masemola were significantly lower compared to those at Mokopane while the reverse was true for the SPAD value where the values were higher at Masemola. Since both NDVI and the SPAD value are a reflection of the chlorophyll content
(Indradewa et al., 2019), these contrasting results are very unusual and not expected. The measured chlorophyll content did not provide much information to explain the observed contrast in NDVI and the SPAD value since there were no significant differences between the locations until after the chilling damage at 70 DAP (Fig 3).
Leaf area index on the other hand varied significantly at 45, 60 and 75 DAP. Masemola had significantly lower LAI throughout when compared to the two locations. The highest LAI was observed at Dalmada at 45 DAP, but at the later stages of growth, the highest was recorded at Mokopane. The findings suggest that the plant growth was more vigorous at Mokopane which agrees with the higher biomass observed in Fig 2. Therefore, based on the observed grain yield and the biomass at these two locations, it seems that LAI and NDVI showed a better reflection of the yield compared to the SPAD value.
Bivariate correlation between NDVI and SPAD value showed that the two had positive relationships which were only significant at Masemola. The correlation analysis also showed that chlorophyll content was positively related to both but was again only significant at Masemola. In addition, it was observed that biomass and grain yield were negatively related to SPAD and chlorophyll content but showed a strong positive relationship with NDVI,
E and
gs (Table 2). On the other hand, both InsWUE and IntrWUE showed negative relationships with
Ci, E and
gs but were more strongly related to
Ci, with R-values of -0.82 and -0.90 respectively (Table 3). InsWUE did not show any significant relationship with SPAD, NDVI, Chlorophyll and LAI while IntrWUE only showed a significant negative relationship with NDVI.
The effect of DAP on SPAD value, NDVI and LAI was only analysed for Mokopane and Masemola due to missing data at Dalmada. Masemola, differed significantly with other locations hence Fig 4 shows the comparison between Masemola and Mokopane. The SPAD value did not vary much with time from 18 DAP to 60 DAP at each location but still showed differences between the two locations (Fig 4). There was a significant reduction in the SPAD value at 75 DAP which was more abrupt at Masemola. The SPAD value at Masemola plummeted by almost 30 SPAD value units from 55 to 27 compared to a 10 unit drop observed at Mokopane. LAI index on the other generally increased with DAP at both locations until 60 DAP before reducing at 75 DAP. Unlike the SPAD value, LAI was higher at Mokopane compared to at Masemola. The LAI index at Mokopane ranged from 1.8 to as high as 4.8 compared to values of 0.6 to 2.0 observed at Masemola. NDVI showed a similar trend to that of LAI. NDVI values at Mokopane were consistently higher than those at Masemola. Fig 4 shows that NDVI increased with DAP, peaking at 45 DAP before decreasing again. The variation in NDVI of dry bean plants over the season could be described by the quadratic equation on Fig 4.
Seasonal variation in leaf gas exchanges with location and DAP
In addition to the three parameters discussed above, leaf gas exchanges were also measured (Table 1 and Fig 5). Firstly, the effect of location on leaf gas exchanges was assessed at different DAP (Table 1). There was an inclination for intercellular CO
2 concentration to be higher at 45 DAP when compared to at 60 DAP (Table 1). The values were also relatively higher at Mokopane compared to the other two locations. Dalmada had the highest photosynthesis rate (11.65 μmol m
-2 s
-1) at 45 DAP compared to other locations. However, at 60 DAP the highest photosynthesis rate was observed at Mokopane with 9.41 μmol m
-2 s
-1. Stomatal conuctance was higher at Mokopane at both 45 and 60 DAP but Masemola and Dalmada did not show any difference. Transpiration rate was also significantly higher at Mokopane compared to the other locations at 60 DAP. Water use efficiency (WUE) measured at leaf level also showed significant differences between the locations. Both instantaneous and intrinsic WUE were lower at Mokopane compared to Masemola and Dalmada which did not differ between themselves.
Secondly, the effect of DAP on leaf gas exchanges were assessed (Fig 5). Intercellular CO
2 concentration was relatively low at Masemola compared to Mokopane. Also,
A fluctuated more at Mokopane compared to at Masemola. Intercellular CO
2 concentration (
Ci) at Mokopane ranged from 253 to 409 ppm while it ranged from 159 to 281 ppm at Masemola. Despite Masemola having relatively low
Ci, it had higher photosynthesis rate at 18 and 32 DAP which however dramatically declined from 10.38 to 4.48 μmol m
-2 s
-1. Transpiration rate was also lower at Masemola and constantly decreased with time from 18 to 60 DAP while the transpiration rate at Mokopane initially decreased before increasing again at 60 DAP. Stomatal conductance (
gs) at Masemola also declined with time as observed with
E but the
gs at Mokopane fluctuated with DAP.
The fluctuation of
gs at Mokopane can be attributed to variations in soil moisture levels, probably due to some measurements being taken soon after irrigation events while others may have been taken several days after irrigation. Such variations in soil moisture can be mirrored in leaf water status, particularly in isohydric plants
(Igarashi et al., 2015). Many studies have shown that
gs responds to variations in soil moisture level
(Serret et al., 2018, Munjonji et al., 2017). These previous findings suggest that at Masemola, the crop might have been subjected to continuous soil drying thus resulting in continuous decrease in
gs. When roots are subjected to a drying soil they produce abscisic acid which in turn signal the closure of the stomata leading to reduction in
gs, and subsequently
E and
A (Saradadevi
et al., 2014,
Osakabe et al., 2014). This is also supported by the observed data in Fig 5.
The lower rates of
E, gs and
Ci observed at Masemola resulted in higher instantaneous WUE (InsWUE) and intrinsic WUE (IntrWUE) (Fig 6). Initially, reduction in
gs reduces
E more than it reduces
A resulting in higher water use efficiency at leaf level i.e. IntrWUE and InsWUE (Nobel, 2009). In Fig 6, it was observed that both IntrWUE and InsWUE were significantly higher and increased with time (i.e. with DAP) at Masemola compared to Mokopane despite
gs at Masemola always decreasing. However, as the soil continued to dry out and crop water stress became more severe, both
E and
A were reduced drastically due to wilting of the leaves. The results showed that higher WUE at leaf level does not necessarily result in better yields but only proves that crops tend to use water more efficiently under limited water supply compared to when well-watered. Although this is not completely new finding in plant sciences, it however emphasises the need for deficit irrigation in drier areas where water supply is limited and hence the need to efficiently utilise the little water available.