Experimental and computational comparison of freeze-thaw-induced pressure generation in red and sugar maple.

Sap exudation is the process whereby trees such as sugar (Acer saccharum Marsh.) and red maple (Acer rubrum L.) generate unusually high positive stem pressure in response to repeated cycles of freeze and thaw. This elevated xylem pressure permits the sap to be harvested over a period of several weeks and hence is a major factor in the viability of the maple syrup industry. The extensive literature on sap exudation documents competing hypotheses regarding the physical and biological mechanisms that drive positive pressure generation in maple, but to date, relatively little effort has been expended on devising mathematical models for the exudation process. In this paper, we utilize an existing model of Graf et al. (J Roy Soc Interface 12:20150665, 2015) that describes heat and mass transport within the multiphase gas-liquid-ice mixture in the porous xylem tissue. The model captures the inherent multiscale nature of xylem transport by including phase change and osmotic transport in wood cells on the microscale, which is coupled to heat transport through the tree stem on the macroscale. A parametric study based on simulations with synthetic temperature data identifies the model parameters that have greatest impact on stem pressure build-up. Measured daily temperature fluctuations are then used as model inputs and the resulting simulated pressures are compared directly with experimental measurements taken from mature red and sugar maple stems during the sap harvest season. The results demonstrate that our multiscale freeze-thaw model reproduces realistic exudation behavior, thereby providing novel insights into the specific physical mechanisms that dominate positive pressure generation in maple trees.

Quantitative examination of the anatomy of the juvenile sugar maple xylem.

New methodologies have enabled viable sap yields from juvenile sugar maple trees. To further improve yields, a better understanding of sap exudation is required. To achieve this, the anatomy of the xylem must first be fully characterised. We examine juvenile maple saplings using light optical microscopy (LOM) and scanning electron microscopy (SEM), looking at sections cut along differing orientations as well as macerations. From this we measure various cell parameters. We find diameter and length of vessel elements to be 28 ± 8 µm and 200 ± 50 µm, for fibre cells 8 ± 3 µm and 400 ± 100 µm, and for ray parenchyma cells 8 ± 2 µm and 50 ± 20 µm. We also examine pitting present on different cell types. On vessel elements we observe elliptical bordered pits connecting to other vessel elements (with major axis of 2.1 ± 0.7 µm and minor 1.3 ± 0.3 µm) and pits connecting to ray parenchyma (with major axis of 4 ± 2 µm and minor 2.0 ± 0.7 µm). We observe two distinct pit sizes on fibres with circular pits 0.7 ± 0.2 µm in diameter and ellipsoidal pits 1.6 ± 0.4 µm by 1.0 ± 0.3 µm. We do not observe distinct pitting patterns on different fibre types. The various cell and pit measurements obtained generally agree with the limited data available for mature trees, with the exception of vessel element and fibre length, both of which were significantly smaller than reported values.

Examination of embolisms in maple and birch saplings utilising microCT.

We demonstrate the application of synchrotron x-ray microtomography (microCT) to non-invasively examine the internal structure of a maple and birch sapling. We show that, through the use of standard image analysis techniques, embolised vessels can be extracted from reconstructed slices of the stem. By combining these thresholded images with connectivity analysis, we map out the embolisms within the sapling in three dimensions and evaluate the size distribution, showing that large embolisms over 0.005 mm3 in volume compose the majority of the saplings' total embolised volume. Finally we evaluate the radial distribution of embolisms, showing that in maple fewer embolisms are present towards the cambium, while birch has a more uniform distribution.

Maple syrup-production, composition, chemistry, and sensory characteristics.

Maple syrup is made from sap exuded from stems of the genus Acer during the springtime. Sap is a dilute solution of primarily water and sucrose, with varying amounts of amino and organic acids and phenolic substances. When concentrated, usually by heating, a series of complex reactions produce a wide variety of flavor compounds that vary due to processing and other management factors, seasonal changes in sap chemistry, and microbial contamination. Color also forms during thermal evaporation. Flavor and color together are the primary factors determining maple syrup grade, and syrup can range from very light-colored and delicate-flavored to very dark-colored and strong-flavored.

Anthocyanin influence on light absorption within juvenile and senescing sugar maple leaves - do anthocyanins function as photoprotective visible light screens?

Foliar anthocyanins are hypothesised to function as photoprotective visible light screens, preventing over-excitation of the photosynthetic system, and decreasing the likelihood of photo-oxidative stress by absorbing green light and reducing the amount of light available to be absorbed by chloroplasts in deeper tissue layers. Chlorophyll fluorescence imaging was used to test the hypothesis that anthocyanins in the palisade mesophyll of juvenile and senescing sugar maple (Acer saccharum Marsh.) leaves function as visible light screens by assessing their influence on light absorption profiles within leaves. We hypothesised that an effective anthocyanic light screen should reduce light absorption, particularly of green wavelengths, by chloroplasts in the spongy mesophyll. Both anthocyanic juvenile and senescing leaves absorbed greater amounts of green light than corresponding nonanthocyanic leaves. However, profiles of green light absorption by chlorophyll within anthocyanic leaves were not shifted to reflect reduced absorption of green light by spongy mesophyll chloroplasts. Further, the spongy mesophyll of both anthocyanic juvenile and senescing leaves absorbed proportions of green light equal to or greater than the spongy mesophyll of corresponding nonanthocyanic leaves. These results indicate that though they may provide a general source of photoprotection by reducing the total quantity of light available to be absorbed by chlorophyll, the anthocyanins in juvenile and senescing sugar maple leaves do not attenuate light in a manner consistent with that expected for an anthocyanic screen in the palisade mesophyll.

Contribution of anthocyanins to the antioxidant capacity of juvenile and senescing sugar maple (Acer saccharum) leaves.

Foliar anthocyanins are hypothesised to provide an additional source of photoprotection from photooxidative stress to the leaves in which they occur through their ability to scavenge excess free radical species. Although demonstrated to significantly enhance the antioxidant status of red morphs of fully expanded leaves of some species, the contribution of anthocyanins to the antioxidant capacity of the juvenile and senescing leaves in which they frequently occur has not been examined. Antioxidant activity of extracts from anthocyanic and non-anthocyanic juvenile and senescing sugar maple (Acer saccharum Marsh.) leaves from similar light environments was assessed using the stable free radical 1,1-diphenyl-2-picryl hydrazyl (DPPH). Anthocyanin content was significantly correlated with antioxidant activity in extracts of anthocyanic juvenile leaves but only weakly correlated in extracts of anthocyanic senescing leaves. In addition, the antioxidant activity of anthocyanic and non-anthocyanic leaves was equal in both juvenile and senescing leaves. Thus, although anthocyanins may contribute to the antioxidant capacity of anthocyanic juvenile and senescing sugar maple leaves, these results are not consistent with the hypothesis that anthocyanins provide an enhancement to the photoprotection available in either leaf type through free radical scavenging. The results suggest anthocyanins may be part of alternative strategies employed by anthocyanic juvenile and senescing maple leaves to achieve similar levels of antioxidant capacity as their non-anthocyanic counterparts to cope with the same set of environmental challenges.