Leaf area (cm²)
Processed poultry manure application significantly affected leaf area (Fig 1). Compared with P0 (20.01 cm
2), P1 increased leaf area to 21.55 cm2, but this difference was not significant). In contrast, P2 significantly increased leaf area to 23.94 cm
2. For CPPU, C1 produced a leaf area of 19.94 cm
2, which did not differ significantly from C0 (19.59 cm
2). However, C2 and C3 significantly increased leaf area to 23.07 and 24.72 cm
2, respectively, compared with C0. The poultry manure x CPPU interaction was also significant (Table 2), with the highest leaf area recorded under P2C3 and the lowest under P0C0.
P0, P1 and P2 represent processed poultry manure application rates of 0, 75 and 150 g pot
-1, respectively. C0, C1, C2 and C3 represent foliar forchlorfenuron (CPPU) concentrations of 0, 3, 6 and 9 mg L
-1, respectively.
The increase in leaf area under P2 may reflect improved nutrient supply and growing-medium conditions following processed poultry manure application. Organic amendments can improve soil physical properties, potentially providing more favorable conditions for root development and nutrient acquisition
(Sruthi et al., 2025; Donbiaksiam et al., 2026). Nitrogen supplied by the manure supports amino acid and protein synthesis, as well as the cellular processes involved in leaf development and expansion
(Wang et al., 2024; Zayed et al., 2023). Potassium may also contribute through its role in osmotic adjustment and the maintenance of cell turgor required for expansion
(Hasanuzzaman et al., 2018). The significant increases under C2 and C3, compared with C0, are consistent with the cytokinin activity of CPPU, which can promote cell division and leaf development
(Werner et al., 2001). The highest value under P2C3 suggests that adequate nutrient supply may support the leaf expansion response to CPPU, although the underlying mechanisms were not measured directly.
Total chlorophyll content of the leaves (mg g-1 FW)
Processed poultry manure significantly affected total chlorophyll content (Fig 2). Both P1 (1.28 mg g
-1 FW) and P2 (1.32 mg g
-1 FW) significantly increased total chlorophyll compared with P0 (1.22 mg g
-1 FW). For CPPU, C1 (1.25 mg g
-1 FW) did not differ significantly from C0 (1.23 mg g
-1 FW), whereas C3 significantly increased total chlorophyll to 1.34 mg g
-1 FW. C2 recorded 1.28 mg g
-1 FW. The poultry manure x CPPU interaction was significant (Table 2), with the highest total chlorophyll content recorded under P2C3 and the lowest under P0C0.
The higher total chlorophyll content under P1 and P2 may be associated with the nutritional contribution of processed poultry manure, particularly its nitrogen and magnesium contents. Nitrogen supports chlorophyll synthesis and the formation of proteins involved in the photosynthetic apparatus (
Fathi, 2022), while magnesium contributes to chlorophyll structure and the activation of enzymes involved in photosynthetic metabolism
(Senbayram et al., 2015). The significant increase under C3 may reflect the influence of CPPU on chloroplast development and the maintenance of leaf chlorophyll, consistent with the physiological functions of cytokinins
(Sakakibara et al., 2006). Cytokinin-related regulation of nucleic acid and protein synthesis may also support these processes
(Smith et al., 2017). The highest chlorophyll content under P2C3 may therefore reflect complementary nutritional and growth-regulatory effects. However, increased chlorophyll content alone does not establish an increase in net photosynthetic rate.
Phosphorus percentage in the leaves
Processed poultry manure significantly affected leaf phosphorus concentration (Fig 3). Both P1 (0.42%) and P2 (0.43%) significantly increased leaf phosphorus compared with P0 (0.37%). Similarly, CPPU application significantly increased leaf phosphorus at C1 (0.41%), C2 (0.42%) and C3 (0.44%) compared with C0 (0.34%). The highest concentrations were recorded under P2 and C3 for the respective factors. However, the poultry manure x CPPU interaction was not significant (Table 2).
The significant increases in leaf phosphorus under P1 and P2 may partly result from phosphorus supplied by processed poultry manure. Organic matter decomposition may additionally modify phosphorus availability through changes in soil chemical conditions and interactions involving humic substances
(Kobierski et al., 2017). Changes in rhizosphere pH could also influence nutrient availability, although their direction and magnitude depend on the amendment and soil conditions (
Raghavendar, 2026). Phosphorus is involved in energy transfer and the formation of compounds essential for plant metabolism and growth
(Khan et al., 2023; Al-Dulaimy and Mohammed, 2024). The higher leaf phosphorus concentrations under all CPPU treatments may be associated with cytokinin-mediated changes in nutrient allocation toward developing tissues (
Davies, 2004). Nevertheless, leaf concentration measurements alone cannot distinguish increased uptake from altered internal distribution. The nonsignificant interaction indicates that a combined effect beyond the individual factor effects was not demonstrated for this trait.
Potassium percentage in the leaves
Processed poultry manure significantly affected leaf potassium concentration (Fig 4). P2 significantly increased leaf potassium to 2.04% compared with 1.89% under P0, while P1 recorded 1.97%. For CPPU, C1 (1.91%) did not differ significantly from C0 (1.84%). However, C2 (1.98%) and C3 (2.15%) significantly increased leaf potassium compared with C0. The poultry manure x CPPU interaction was significant (Table 2), with the highest leaf potassium concentration recorded under P2C3 and the lowest under P0C0.
The significantly higher leaf potassium concentration under P2 may be attributed partly to potassium supplied by processed poultry manure and its potential effects on nutrient availability in the growing medium (
Al-Jaf et al., 2023). Potassium supports enzyme activation, photosynthesis and carbohydrate metabolism
(Rawat et al., 2022). The significant increases under C2 and C3 may reflect changes in nutrient demand and allocation associated with CPPU-induced leaf development. Greater leaf area and chlorophyll content could potentially support carbon assimilation and the metabolic processes associated with nutrient acquisition, although these processes were not directly assessed (
Taiz and Zeiger, 2010). The highest potassium concentration under P2C3 suggests that the response to CPPU depended partly on manure supply. This interpretation is broadly consistent with the nutritional responses to CPPU-containing treatments reported by
Mohammed and Al-Dulaimy (2023) and
Al-Hameedi and Al-Tamimi (2025).
Calcium percentage in the leaves
Processed poultry manure significantly affected leaf calcium concentration (Fig 5). P1 (1.29%) did not differ significantly from P0 (1.27%), whereas P2 significantly increased leaf calcium to 1.36%. For CPPU, C1 (1.27%) did not differ significantly from C0 (1.26%), while C3 significantly increased leaf calcium to 1.38%. C2 recorded a calcium concentration of 1.32%. The poultry manure x CPPU interaction was significant (Table 2), with the highest leaf calcium concentration recorded under P2C3 and the lowest under P0C0.
The significant increase in leaf calcium concentration under P2 may partly reflect the calcium supplied by processed poultry manure. Calcium contributes to cell-wall structure through its association with pectic substances and supports tissue integrity during development
(Kour et al., 2023). It also plays important roles in membrane stability and cellular signaling (
Thor, 2019), as well as the regulation of cellular processes associated with growth
(Wdowiak et al., 2024). The higher calcium concentration under C3 may reflect indirect effects of CPPU on leaf development and calcium delivery to growing tissues. However, the present measurements do not establish whether this response resulted from altered uptake, transport or tissue growth. The highest concentration under P2C3 suggests that manure supply influenced the calcium response to CPPU, consistent with the significant interaction.
Magnesium percentage in the Leaves
Processed poultry manure significantly affected leaf magnesium concentration (Fig 6). P1 (0.44%) did not differ significantly from P0 (0.42%), whereas P2 significantly increased leaf magnesium to 0.51%. For CPPU, C1 (0.45%) did not differ significantly from C0 (0.42%), while C3 significantly increased leaf magnesium to 0.50%. C2 recorded a magnesium concentration of 0.47%. The poultry manure x CPPU interaction was significant (Table 2), with the highest leaf magnesium concentration recorded under P2C2 and the lowest under P0C0.
The significantly higher leaf magnesium concentration under P2 may partly result from the magnesium supplied by processed poultry manure. Magnesium supports photosynthesis and numerous enzyme reactions involved in energy metabolism
(Senbayram et al., 2015). It also contributes to carbohydrate transport from source leaves to sink organs
(Ferreira et al., 2023) and participates in physiological responses to abiotic stress
(Ahmed et al., 2023), although stress tolerance was not evaluated in this experiment
(Walaa et al., 2026). The significant increase under C3 may be associated with CPPU-related changes in leaf development and nutrient allocation. The highest magnesium concentration under P2C2, despite the highest CPPU main-effect mean occurring under C3, illustrates the importance of considering treatment combinations when interpreting the significant interaction. The beneficial nutritional response to manure application is broadly consistent with the olive studies cited by
Khalil and Abdulqader (2023) and
Almashhadani and Al-Hadethi (2025).