TL;DR: Recognizing the diversity of belowground bud banks is the starting point for understanding the many evolutionary pathways available for responding to severe recurrent disturbances.
Abstract: Despite long-time awareness of the importance of the location of buds in plant biology, research on belowground bud banks has been scant. Terms such as lignotuber, xylopodium and sobole, all referring to belowground bud-bearing structures, are used inconsistently in the literature. Because soil efficiently insulates meristems from the heat of fire, concealing buds below ground provides fitness benefits in fire-prone ecosystems. Thus, in these ecosystems, there is a remarkable diversity of bud-bearing structures. There are at least six locations where belowground buds are stored: roots, root crown, rhizomes, woody burls, fleshy swellings and belowground caudexes. These support many morphologically distinct organs. Given their history and function, these organs may be divided into three groups: those that originated in the early history of plants and that currently are widespread (bud-bearing roots and root crowns); those that also originated early and have spread mainly among ferns and monocots (nonwoody rhizomes and a wide range of fleshy underground swellings); and those that originated later in history and are strictly tied to fire-prone ecosystems (woody rhizomes, lignotubers and xylopodia). Recognizing the diversity of belowground bud banks is the starting point for understanding the many evolutionary pathways available for responding to severe recurrent disturbances.
TL;DR: It is found that plant mortality after multiple clipping was 10% for Arbutus and 30% for Erica, and was primarily attributed to exhaustion of carbon reserves because starch concentrations decreased by 96% in dead plants.
Abstract: 1. We investigated whether reserves stored in the lignotubers of two Mediterranean shrubs, Arbutus unedo and Erica arborea, were significantly mobilized to support the demands of regrowth and respiration after clipping the tops at different frequencies.
2. After a single clipping, Arbutus showed a 29% decrease of phosphorus concentration by the end of the first growing season. Two years after recovery from clipping, the starch levels remained lower than those of unclipped plants. Similarly, Erica showed depletion of starch, but no nutrient reserves were depleted significantly.
3. Regrowth after multiple clippings mobilized a large fraction of the starch and nutrients stored in the lignotuber. Mean starch concentrations were depleted by 87–93% after multiple clippings and concentrations of nitrogen, phosphorus, potassium and magnesium were depleted by 10–45%, 27–41%, 19–39% and 23–31%, respectively.
4. An average-sized lignotuber produced 288 resprouts for Arbutus and 1990 resprouts for Erica during a 27 month period of multiple clippings, at the end of which the first plants died.
5. Plant mortality after multiple clipping was 10% for Arbutus and 30% for Erica, and was primarily attributed to exhaustion of carbon reserves because starch concentrations decreased by 96% in dead plants.
TL;DR: Drought appears to interact with fire by altering postfire shrub recovery and altering species abundances and composition of chaparral communities.
Abstract: We examined postfire regeneration of chaparral shrubs during an intense drought. This study focused on the demography and physiology of shrub species that resprout from a basal lignotuber following fire. We found significant levels of resprout mortality when intense drought occurred in the year following fire during the period of shrub recovery. Three of the seven sampled resprouting species had the greatest or near greatest levels of mortality ever recorded when compared to previous studies. Most shrub mortality occurred during the drought after individuals had resprouted (i.e. individuals survived fire, resprouted and then subsequently died). Physiological measurements of species with high mortality suggested that resprout stems were highly embolized and xylem hydraulic conductivities were close to zero during the peak of the drought. In addition, lignotubers of two of the three species experiencing high mortality were depleted of starch. Population densities of most shrub species declined after the drought compared with their prefire levels, with the exception of one drought tolerant obligate seeding species. Resprouting shrub species may deplete their carbohydrate reserves during the resprouting process, making them particularly vulnerable to drought because of the need to transpire water to acquire the CO2 that is used to supply energy to a large respiring root system. Drought appears to interact with fire by altering postfire shrub recovery and altering species abundances and composition of chaparral communities.
TL;DR: The results suggest that stabilizing processes, probably at the stand level, may have been operating in the long-term to mitigate any impact of drought on NSC levels, and highlight the necessity to incorporate long- term experimental studies of plant responses to drought.
Abstract: Stored non-structural carbohydrates (NSC) have been proposed as a key determinant of drought resistance in plants. However, the evidence for this role is controversial, as it comes mostly from observational, short-term studies. Here, we take advantage of a long-term experimental throughfall reduction to elucidate the response of NSC to increased drought 14 years after the beginning of the treatment in three Mediterranean woody species (Quercus ilex L., Arbutus unedo L. and Phillyrea latifolia L.). In addition, we selected 20 Q. ilex individuals outside the experimental plots to directly assess the relationship between defoliation and NSC at the individual level. We measured the seasonal course of NSC concentrations in leaves, branches and lignotuber in late winter, late spring, summer and autumn 2012. Total concentrations of NSC were highest in the lignotuber for all species. In the long-term drought experiment we found significant depletion in concentrations of total NSC in treatment plots only in the lignotuber of A. unedo. At the same time, A. unedo was the only species showing a significant reduction in BAI under the drought treatment during the 14 years of the experiment. By contrast, Q. ilex just reduced stem growth only during the first 4 years of treatment and P. latifolia remained unaffected over the whole study period. However, we found a clear association between the concentrations of NSC and defoliation in Q. ilex individuals sampled outside the experimental plots, with lower total concentrations of NSC and lower proportion of starch in defoliated individuals. Taken together, our results suggest that stabilizing processes, probably at the stand level, may have been operating in the long-term to mitigate any impact of drought on NSC levels, and highlight the necessity to incorporate long-term experimental studies of plant responses to drought.
TL;DR: Pronounced swelling of the upper root region suggests ontogenetic development of a lignotuber in the resprouter forms of the two Erica species, and the distinct allocation of starch in roots seems to be genetically determined.
Abstract: Post-fire sprouting of dormant buds in resprouter plants is facilitated by stored carbohydrate reserves, with starch being the critical reserve. Starch is mainly stored in xylem parenchyma ray tissue of woody underground organs, such as burls, lignotubers, and roots. We carried out a comparative analysis of the pattern of starch storage and the proportion of parenchymatic ray tissue in the upper root or cotyledonary region of seedlings from seeder and resprouter forms within two Cape Erica (Ericaceae) species: E. coccinea L. and E. calycina L., which were raised in the greenhouse under controlled irrigation. We also explored the root-to-shoot allocation patterns of seeder and resprouter seedlings in these two species. Resprouter seedlings of both species showed higher relative amounts of upper-root starch and upper-root storage tissue as well as a higher root-to-shoot allocation than their seeder counterparts. Pronounced swelling of the upper root region suggests ontogenetic development of a lignotuber in the resprouter forms of the two Erica species. The distinct allocation of starch in roots seems to be genetically determined and would account for the apparent differences in the root-to-shoot allocation patterns between both regeneration forms from the early seedling stage.