Effects of phloem on canopy dieback, tested with manipulations and a canker pathogen in the Corylus avellana/Anisogramma anomala host/pathogen system.

Canker pathogens cause necrosis of the phloem, but in many host/pathogen systems, they also cause canopy dieback, which implicates xylem, not phloem dysfunction. We hypothesize that this dieback distal to the canker is caused by water stress resulting from the lack of a phloem-to-xylem connection, which in a healthy plant would allow delivery of nonstructural carbohydrates (NSCs) and water inward to aid in xylem embolism refilling. We tested several components of this hypothesis in the host/pathogen system Corylus avellana L./Anisogramma anomala (Peck) E. Müll (Eastern filbert blight). Cankers were non-girdling and usually ≥0.1 m long. As expected, healthy controls had higher specific conductivity (Ks) than diseased stems, but unexpectedly, had similar moisture content (m.c.), showing that the lower Ks did not result from more embolisms in the diseased stems. Moreover, manipulations that removed cambium and phloem to simulate a canker, or that shaded stems to lower NSCs, did not result in lower Ks or m.c. than controls. The outer millimeter of xylem adjacent to a canker had infrequent tyloses and/or fungal hyphae in many but not all samples, and dye studies showed little xylem water transport in that region, but the incidence of these blockages was insufficient to cause the observed 19% decrease in Ks. Healthy stems had higher m.c. than diseased stems above the canker (or analogous) location and were longer for the same leaf weight, suggestive of water stress in the upper portion of diseased stems. These results suggest that dieback distal to cankers in this system results from the bottleneck in water transport in the region adjacent to a canker, but did not find evidence to support the requirement of a phloem-to-xylem connection for continued water transport.

Coordination of leaf structure and gas exchange along a height gradient in a tall conifer.

The gravitational component of water potential and frictional resistance during transpiration lead to substantial reductions in leaf water potential (Psi(l)) near the tops of tall trees, which can influence both leaf growth and physiology. We examined the relationships between morphological features and gas exchange in foliage collected near the tops of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees of different height classes ranging from 5 to 55 m. This sampling allowed us to investigate the effects of tree height on leaf structural characteristics in the absence of potentially confounding factors such as irradiance, temperature, relative humidity and branch length. The use of cut foliage for measurement of intrinsic gas-exchange characteristics allowed identification of height-related trends without the immediate influences of path length and gravity. Stomatal density, needle length, needle width and needle area declined with increasing tree height by 0.70 mm(-2) m(-1), 0.20 mm m(-1), 5.9 x 10(-3) mm m(-1) and 0.012 mm(2) m(-1), respectively. Needle thickness and mesophyll thickness increased with tree height by 4.8 x 10(-2) mm m(-1) and 0.74 microm m(-1), respectively. Mesophyll conductance (g(m)) and CO(2) assimilation in ambient [CO(2)] (A(amb)) decreased by 1.1 mmol m(-2) s(-1) per m and 0.082 micromol m(-2) s(-1) per m increase in height, respectively. Mean reductions in g(m) and A(amb) of foliage from 5 to 55 m were 47% and 42%, respectively. The observed trend in A(amb) was associated with g(m) and several leaf anatomic characteristics that are likely to be determined by the prevailing vertical tension gradient during foliar development. A linear increase in foliar delta(13)C values with height (0.042 per thousand m(-1)) implied that relative stomatal and mesophyll limitations of photosynthesis in intact shoots increased with height. These data suggest that increasing height leads to both fixed structural constraints on leaf gas exchange and dynamic constraints related to prevailing stomatal behavior.

Height-related trends in leaf xylem anatomy and shoot hydraulic characteristics in a tall conifer: safety versus efficiency in water transport.

Hydraulic vulnerability of Douglas-fir (Pseudotsuga menziesii) branchlets decreases with height, allowing shoots at greater height to maintain hydraulic conductance (K shoot) at more negative leaf water potentials (Psi l). To determine the basis for this trend shoot hydraulic and tracheid anatomical properties of foliage from the tops of Douglas-fir trees were analysed along a height gradient from 5 to 55 m. Values of Psi l at which K shoot was substantially reduced, declined with height by 0.012 Mpa m(-1). Maximum K shoot was reduced by 0.082 mmol m(-2) MPa(-1) s(-1) for every 1 m increase in height. Total tracheid lumen area per needle cross-section, hydraulic mean diameter of leaf tracheid lumens, total number of tracheids per needle cross-section and leaf tracheid length decreased with height by 18.4 microm(2) m(-1), 0.029 microm m(-1), 0.42 m(-1) and 5.3 microm m(-1), respectively. Tracheid thickness-to-span ratio (tw/b)2 increased with height by 1.04 x 10(-3) m(-1) and pit number per tracheid decreased with height by 0.07 m(-1). Leaf anatomical adjustments that enhanced the ability to cope with vertical gradients of increasing xylem tension were attained at the expense of reduced water transport capacity and efficiency, possibly contributing to height-related decline in growth of Douglas fir.

Phytophthora ramorum Colonizes Tanoak Xylem and Is Associated with Reduced Stem Water Transport.

ABSTRACT Isolation, detection with diagnostic polymerase chain reaction (PCR), and microscopy demonstrated the presence of Phytophthora ramorum in the sapwood of mature, naturally infected tanoak (Lithocarpus densiflorus) trees. The pathogen was strongly associated with discolored sapwood (P < 0.001), and was recovered or detected from 83% of discolored sapwood tissue samples. Hyphae were abundant in the xylem vessels, ray parenchyma, and fiber tracheids. Chlamydospores were observed in the vessels. Studies of log inoculation indicated that P. ramorum readily colonized sapwood from inoculum placed in the bark, cambium, or sapwood. After 8 weeks, radial spread of P. ramorum in sapwood averaged 3.0 to 3.3 cm and axial spread averaged 12.4 to 18.8 cm. A field study was conducted to determine if trees with infected xylem had reduced sap flux and reduced specific conductivity relative to noninfected control trees. Sap flux was monitored with heat-diffusion sensors and tissue samples near the sensors were subsequently tested for P. ramorum. Adjacent wood sections were excised and specific conductivity measured. Both midday sap flux and specific conductivity were significantly reduced in infected trees versus noninfected control trees. Vessel diameter distributions did not differ significantly among the two treatments, but tyloses were more abundant in infected than in noninfected trees. Implications for pathogenesis, symptomology, and epidemiology are discussed.