Carbohydrate and Nutritional Responses to Stem Girdling and Drought Stress with Respect to Understanding Symptoms of Huanglongbing in Citrus

The most important worldwide problem in citrus production is the bacterial disease Huanglongbing (HLB; citrus greening) caused by a phloem-limited bacterium Candidatus Liberibacter asiaticus. The earliest visible symptoms of HLB on leaves are vein yellowing and an asymmetrical chlorosis referred to as blotchy mottle, thought to be the result of starch accumulation. We tested the hypothesis that such visible symptoms are not unique to HLB by stem girdling 2-year-old seedlings of two citrus rootstocks with and without drought stress in the greenhouse. After 31 days, girdling had little effect on shoot growth but girdling increased the relative growth rate of shoots in drought-stressed trees. Starch content in woody roots of non-girdled trees was three to 19 times higher than in girdled trees. In non-girdled trees, drought stress induced some starch accumulation in roots, but there were no effects of drought stress on root starch or sucrose in girdled trees. Girdling induced a 4-fold greater starch content in leaves on well-watered trees but leaf sucrose content was unaffected. Girdling reduced leaf transpiration in well-watered trees but net assimilation of CO2 was unaffected by girdling or leaf starch accumulation. Leaves on girdled trees clearly had visible blotchy mottle symptoms but no symptoms developed on non-girdled trees. The increase in leaf starch, up to 50% dry weight (DW), resulted in an increase in leaf DW per leaf area (LA) and an artificial reduction of many leaf nutrients on a DW basis. Most of these differences disappeared when expressed on a LA basis. Leaf boron (B), however, was inversely related to leaf starch when both were expressed on a LA basis. In the absence of HLB, girdling increased leaf starch, decreased root starch, and duplicated the asymmetric blotchy mottled visual leaf symptoms that have been associated with HLB-infected trees. This supports our contention that such symptoms generally attributed to HLB are not uniquely related to HLB infection, but rather are directly related to starch accumulation and secondarily to nutrient deficiencies in leaves.