ABSTRACT
The ascomycetous fungus Ophiostoma novo-ulmi is the causative agent of the current Dutch elm disease (DED) pandemic, which has ravaged many tens of millions of European and North American elm trees. Host responses in vascular traits were studied in two Dutch elm hybrids, 'Groeneveld' and 'Dodoens', which show different vascular architecture in the secondary xylem and possess contrasting tolerances to DED. 'Groeneveld' trees, sensitive to DED, possessed a high number of small earlywood vessels. However, these trees showed a poor response to DED infection for the earlywood vascular characteristics. Following infection, the proportion of least vessels with a vessel lumen area less than 2500 µm2 decreased from 65.4% down to 53.2%. A delayed response in the increasing density of vessels showing a reduced size in the latewood prevented neither the rapid fungal spread nor the massive colonisation of the secondary xylem tissues resulting in the death of the infected trees. 'Dodoens' trees, tolerant to DED, possessed a low number of large earlywood vessels and showed a prominent and fast response to DED infection. Vessel lumen areas of newly formed earlywood vessels were severely reduced together with the vessel size : number ratio. Following infection, the proportion of least vessels with a vessel lumen area less than 2500 µm2 increased from 75.6% up to 92.9%. A trend in the increasing density of vessels showing a reduced size was maintained not only in the latewood that was formed in the year of infection but also in the earlywood that was formed in the consecutive year. The occurrence of fungal hyphae in the earlywood vessels that were formed a year following the infection was severely restricted, as revealed by X-ray micro-computed tomography imaging. Possible reasons responsible for a contrasting survival of 'Groeneveld' and 'Dodoens' trees are discussed.
ABSTRACT
BACKGROUND AND AIMS: Previous studies have shown that Ophiostoma novo-ulmi, the causative agent of Dutch elm disease (DED), is able to colonize remote areas in infected plants of Ulmus such as the leaf midrib and secondary veins. The objective of this study was to compare the performances in leaf traits between two Dutch elm hybrids 'Groeneveld' and 'Dodoens' which possess a contrasting tolerance to DED. Trait linkages were also tested with leaf mass per area (LMA) and with the reduced Young's modulus of elasticity (MOE) as a result of structural, developmental or functional linkages. METHODS: Measurements and comparisons were made of leaf growth traits, primary xylem density components, gas exchange variables and chlorophyll a fluorescence yields between mature plants of 'Groeneveld' and 'Dodoens' grown under field conditions. A recently developed atomic force microscopy technique, PeakForce quantitative nanomechanical mapping, was used to reveal nanomechanical properties of the cell walls of tracheary elements such as MOE, adhesion and dissipation. KEY RESULTS: 'Dodoens' had significantly higher values for LMA, leaf tissue thickness variables, tracheary element lumen area (A), relative hydraulic conductivity (RC), gas exchange variables and chlorophyll a fluorescence yields. 'Groeneveld' had stiffer cell walls of tracheary elements, and higher values for water-use efficiency and leaf water potential. Leaves with a large carbon and nutrient investment in LMA tended to have a greater leaf thickness and a higher net photosynthetic rate, but LMA was independent of RC. Significant linkages were also found between the MOE and some vascular traits such as RC, A and the number of tracheary elements per unit area. CONCLUSIONS: Strong dissimilarities in leaf trait performances were observed between the examined Dutch elm hybrids. Both hybrids were clearly separated from each other in the multivariate leaf trait space. Leaf growth, vascular and gas exchange traits in the infected plants of 'Dodoens' were unaffected by the DED fungus. 'Dodoens' proved to be a valuable elm germplasm for further breeding strategies.
