Essential for fertility (male and female gametophyte functions and development) (PubMed:28585692, PubMed:29424430). Required for the integrity of female gametic mitochondria (By similarity). Involved in embryo apical-basal patterning, and particularly dorsal-ventral patterning, during early embryogenesis, and endosperm free nucleus positioning and development as well as early endosperm development, probably by modulating the expression pattern of related genes (e.g. AL1, MYB3/AL2, CYP78A13/GE, PNH1, HAZ1, MPK6 and OSH1) (PubMed:28585692). Has function in triggering of endosperm programmed cell death (PCD) leading to syncytial endosperm cellularization and starchy endosperm cell maturation (PubMed:28585692). Implicated in central vacuole dynamics necessary for microspore development leading to pollen production, and for pollen development and germination (PubMed:28585692, PubMed:29424430). Expressed in roots, stems, leaves and florets. In florets, detected in glumes, lemmas, paleas and anthers (PubMed:28585692). Also present in ovaries at various developmental stages (PubMed:28585692). Expressed in the mature embryo sac, which comprises four cell types: egg cell, central cell, synergids, and antipodal cells (PubMed:28585692). Observed throughout early embryogenesis and in the endosperm at both the free-nuclear and cellular endosperm stages (PubMed:28585692). In endosperm, confined progressively to aleurone layer (PubMed:28585692). Expressed in anther tapetum and microspores, as well as in the stigma of mature pistils (PubMed:28585692). In 12 days old seedling roots, detected mainly in the cortex and epidermis of the maturation zone, as well as in root hairs (PubMed:28585692). In stems, concentrated in the pith tissue in the middle of the stem (PubMed:28585692). In leaves, confined to mesophyll tissue (PubMed:28585692). Normal vegetative and floral organ development, but several defects in reproductive organs such as reduced pollen production due to abnormal microspore development, aborted seeds, reduced endosperm nuclei divisions, abnormal distribution of free nuclei missing an oriented positioning at the micropylar end of embryo sacs, and interrupted aleurone differentiation associated with unusual positional endosperm cell differentiation (PubMed:28585692). Disturbed embryos develop slowly, and they exhibit altered pattern formation, particularly the dorsal-ventral pattern and symmetry establishment, thus showing various morphological and structural dysplasias; these phenotypes are associated with missexpression pattern of related genes including derepression of CYP78A13/GE in the scutellar epithelium in an irregular and diffuse pattern, enhanced accumulation of MYB3/AL2, abnormal distribution of AL1 in both endosperm dorsal and ventral sites in the aleurone layer, but reduced levels of OSH1 in embryo indeterminate cells around the presumptive shoot apical meristem (SAM), decreased HAZ1 expression in the outermost cells during early embryogenesis, less MPK6 accumulation and lower levels of PNH1 in the L3 protoderm layer (PubMed:28585692). Impaired triggering of endosperm programmed cell death (PCD) (PubMed:28585692). Altered the central vacuole dynamics leading to disturbed pollen maturation and germination, thus ending in male sterility (PubMed:28585692, PubMed:29424430).