We have cloned and sequenced the nuclear gene of the chloroplast ribosomal protein L21 (rpI2l) of Spinacia oleracea. The gene consists of five exons and four introns. All introns are located in the sequence which corresponds to the Escherichia coli-like central core of the protein. L21 mRNA is present in photosynthetic (leaves) and nonphotosynthetic (roots and seeds) plant organs, although large quantitative differences exist. Primer extension and Si nuclease mapping experiments revealed the existence of two types of transcripts in leaves. The two corresponding start sites were defined as P1 and P2. In roots and seeds, we found only the shorter of the two transcripts (initiated at P2). The nucleotide sequence surrounding P2 resembles promoters for housekeeping and vertebrate r-protein genes. Analysis of several promoter constructions by transient expression confirmed that both transcripts originate from transcription initiation. Results are interpreted to mean that the expression of the rpI21 gene is regulated by alternative promoters. One of the promoters (P2) is constitutive, and the other one (P1) is specifically induced in leaves, i.e., its activation should be related to the transformation of amyloplasts or proplastids to chloroplasts. The gene thus represents the first example of a housekeeping gene which is regulated by the organ-specific usage of alternative promoters. Primer extension analysis and S1 nuclease mapping of another nucleus-encoded chloroplast ribosomal protein gene (rpsl) give evidence that the same type of regulation by two-promoter usage might be a more general phenomenon of plant chloroplast-related ribosomal protein genes. Preliminary results indicate the presence of conserved sequences within the rpl2l and rpsl promoter regions which compete for the same DNA binding activities.Plastids are essential components of plant cells. Depending on their function within a defined plant tissue or organ, plastids differentiate into several types of organelles. They are present as chloroplasts in photosynthetically active cells of leaves, as amyloplasts and proplastids in root cells and seeds, and as chromoplasts in fruit and flower cells.The transformation of plastids into these specialized types of organelles is necessarily based on changes in gene expression within the organelles as well as changes in the expression of nuclear-encoded plastid proteins. These changes concern the induction and/or repression of the expression of plastid-specific genes, but they also have to include modulated expression of genes which are necessary for the genetic functions of all types of organelles. In this respect, we are especially interested in how different types of plastids maintain their translational capacities, e.g., in the regulation of ribosomal gene expression during plastid differentiation.The present knowledge on ribosomal gene expression originates mainly from studies of procaryotic and animal systems. It has been shown previously that the rate of ribosome formation is adapted to the individual cell...