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Pinus densiflora  Siebold et Zucc. is the major green canopy species in the mountainous area of Korea. To assess the response of resin acid biosynthetic genes to mechanical and chemical stimuli, we cloned cDNAs of genes encoding enzymes involved in the 2-C-methyl- d -erythritol 4-phosphate (MEP) pathway (1-deoxy- d -xylulose 5-phosphate synthase (PdDXS), 1-deoxy- d- xylulose 5-phosphate reductoisomerase (PdDXR) and 1-hydroxy-2-methyl-2-( E )-butenyl 4-diphosphate reductase (PdHDR)) by the rapid amplification of cDNA ends (RACE) technique. In addition, we cloned the gene encoding abietadiene synthase (PdABS) as a marker for the site of pine resin biosynthesis.  PdHDR  and  PdDXS  occurred as two gene families. In the phylogenetic trees, PdDXSs, PdDXR and PdHDRs each formed a separate clade from their respective angiosperm homologs.  PdDXS 2,  PdHDR 2 and  PdDXR  were most actively transcribed in stem wood, whereas  PdABS  was specifically transcribed .  The abundance of  PdDXS 2 transcripts in wood in the resting state was generally 50-fold higher than the abundance of  PdDXS 1 transcripts, and  PdHDR 2 transcripts were more abundant by an order of magnitude in wood than in other tissues, with the ratio of  PdHDR 2 to  PdHDR 1 transcripts in wood being about 1. Application of 1 mM methyl jasmonate (MeJA) selectively enhanced the transcript levels of  PdDXS 2 and  PdHDR 2 in wood. The ratios of  PdDXS 2 to  PdDXS 1 and  PdHDR 2 to  PdHDR 1 reached 900 and 20, respectively, on the second day after MeJA treatment, whereas the transcript level of  PdABS  increased twofold by 3 days after MeJA treatment. Wounding of the stem differentially enhanced the transcript ratios of  PdDXS 2 to  PdDXS 1 and  PdHDR 2 to  PdHDR 1 to 300 and 70, respectively. The increase in the transcript levels of the MEP pathway genes in response to wounding was accompanied by two orders of magnitude increase in  PdABS  transcripts. These observations indicated that resin acid biosynthesis activity, represented by  PdABS  transcription, was correlated with the selective transcriptions of  PdDXS 2 and  PdHDR 2. Introduction of  PdDXS 2,  PdHDR 1 and  PdHDR 2 rescued their respective knockout  Escherichia coli  mutants, confirming that at least these three genes were functionally active. Intracellular targeting of the green fluorescent protein fused to the N-terminal 100 amino acid residues of these genes in the  Arabidopsis  transient expression system showed that the proteins were all targeted to the chloroplasts. Our results suggest that the MEP pathway regulates resin biosynthesis in the wood of  P. densiflora  by differential transcription of the multiple  PdDXS a nd  PdHDR  genes.

tree physiology

Regulation of resin acid synthesis in  Pinus densiflora  by differential transcription of genes encoding multiple 1-deoxy- d -xylulose 5-phosphate synthase and 1-hydroxy-2-methyl-2-( E )-butenyl 4-diphosphate reductase genes