Ronald L. Hendrick

The Demography of Fine Roots in a Northern Hardwood Forest

The production, development, and mortality of fine roots in a northern hardwood forest was monitored for 1 yr using minirhizotrons. Roots were divided into two strata based upon their depth in the soil, 30 cm. Cohort analyses of roots produced in the spring of 1989 revealed that while almost 50% of fine roots at both depths survived after 346 d, the number of white roots in each cohort declined very rapidly. Virtually all roots had turned brown after 346 d. The probability of a surviving white root turning brown was much greater than the probability that it would die at all times of the year, and the bulk of root mortality was accounted for by brown roots. Analysis of root length production and mortality showed that total annual length mortality at the 30 cm depth. Fine root production and mortality occurred simultaneously throughout the year, and production was slightly greater than mortality at both depths. Total root length peaked in the summer at both depths, and overwinter production and mortality was rather low. Production of white and brown root length indicated that roots near the soil surface were undergoing much more rapid rates of browning than deep roots. Loss of root length between sampling dates was largely due to roots that died and rapidly decayed or otherwise disappeared.

COUPLING FINE ROOT DYNAMICS WITH ECOSYSTEM CARBON CYCLING IN BLACK SPRUCE FORESTS OF INTERIOR ALASKA

Fine root processes play a prominent role in the carbon and nutrient cycling of boreal ecosystems due to the high proportion of biomass allocated belowground and the rapid decomposition of fine roots relative to aboveground tissues. To examine these issues in detail, major components of ecosystem carbon flux were studied in three mature black spruce forests in interior Alaska, where fine root production, respiration, mortality and decomposition, and aboveground production of trees, shrubs, and mosses were measured relative to soil CO2 fluxes. Fine root production, measured over a two-year period using minirhizotrons, varied from 0.004 ± 0.001 mm·cm–2·d–1 over winter, to 0.051 ± 0.015 mm·cm–2·d–1 during July, with peak growing season values comparable to those reported for many temperate forests using similar methods. On average, 84% of this production occurred within 20 cm of the moss surface, although the proportion occurring in deeper profiles increased as soils gradually warmed throughout the summer. M...

Temporal and depth-related patterns of fine root dynamics in northern hardwood forests

1 The dynamics of fine (< 2.0 mm) roots growing in two northern hardwood forests were quantified to a depth of 1 m in 10-cm increments. We hypothesized that patterns of root production and mortality would be broadly synchronous at all depths, but that deep roots would be comparatively less dynamic than shallow roots. 2 Our data showed that shallow roots were responsible for the majority of total annual fine root production and mortality. Nearly half of all roots growing and dying in the 1 -m profile occurred in the upper 20 cm of the soil, while roots located at depths of 75 cm or more accounted for only 11% of annual production and 4% of annual mortality. 3 Fine root production prior to, or coincident with, canopy expansion was significant at all depths. The relative importance of early fine root growth generally increased with depth, with between 50 and 80% of annual production occurring prior to midJune at depths exceeding 50 cm. Episodic deep root production during the growing season appeared to be related to periods of high water demand. 4 Patterns of fine root mortality were more variable among depths. Mortality was distributed rather evenly throughout the year near the soil surface, but mid-season mortality was generally low at depths greater than 50 cm.

Root respiration in North American forests: effects of nitrogen concentration and temperature across biomes

Root respiration rates have been shown to be correlated with temperature and root N concentration in studies of individual forest types or species, but it is not known how universal these relationships are across forest species adapted to widely different climatic and edaphic conditions. In order to test for broad, cross-species relationships, we measured fine root respiration, as O2 consumption, over a range of temperatures on excised root samples from ten forested study sites across North America in 1997. Significant differences existed among study sites in root respiration rates, with patterns among sites in respiration rate at a given temperature corresponding to differences among sites in fine root N concentrations. Root respiration rates were highly correlated with root N concentrations at all measurement temperatures (r2>0.81, P<0.001, for 6, 18 and 24°C). Lower root respiration rates in gymnosperms than in angiosperms were largely explained by lower fine root N concentrations in gymnosperms, and root N concentrations and respiration rates (at a given temperature) tended to be lower at warm sites (New Mexico, Florida, and Georgia) than at cool sites with short growing seasons (Michigan and Alaska). Root respiration rates increased exponentially with temperature at all sites. The Q10 for root respiration ranged from 2.4 to 3.1, but there were no significant differences among the forest types. The average Q10s for gymnosperms (Q10=2.7) and angiosperms (Q10=2.6) were almost identical, as were the average Q10s for roots of ectomycorrhizal species (Q10=2.7) and arbuscular mycorrhizal species (Q10=2.6). In 1998, fine root respiration at the study sites was measured in the field as CO2 production at ambient soil temperature. Respiration rates under field conditions were dependent on both ambient soil temperature and root N concentration. Relationships between respiration (adjusted for temperature) and root N concentration for the field measurements were similar to those observed in the 1997 laboratory experiments. For root respiration in tree species, it appears that basic relationships with temperature and nitrogen exist across species and biomes.

Relationships among root branch order, carbon, and nitrogen in four temperate species

Abstract The objective of this study was to examine how root length, diameter, specific root length, and root carbon and nitrogen concentrations were related to root branching patterns. The branching root systems of two temperate tree species, Acer saccharum Marsh. and Fraxinus americana L., and two perennial herbs from horizontal rhizomes, Hydrophyllum canadense L. and Viola pubescens Ait., were quantified by dissecting entire root systems collected from the understory of an A. saccharum-Fagus grandifolia Ehrh. forest. The root systems of each species grew according to a simple branching process, with laterals emerging from the main roots some distance behind the tip. Root systems normally consisted of only 4–6 branches (orders). Root diameter, length, and number of branches declined with increasing order and there were significant differences among species. Specific root length increased with order in all species. Nitrogen concentration increased with order in the trees, but remained constant in the perennial herbs. More than 75% of the cumulative root length of tree seedling root systems was accounted for by short (2–10 mm) lateral roots almost always <0.3 mm in diameter. Simple assumptions suggest that many tree roots normally considered part of the dynamic fine-root pool (e.g., all roots <2.0 mm in diameter) are too large to exhibit rapid rates of production and mortality. The smallest tree roots may be the least expensive to construct but the most expensive to maintain based on an increase in N concentration with order.

REGULATION OF FINE ROOT DYNAMICS BY MAMMALIAN BROWSERS IN EARLY SUCCESSIONAL ALASKAN TAIGA FORESTS

The effects of browsing by moose and snowshoe hares on fine root production, mortality, and decomposition in early successional forest ecosystems along the Tanana River floodplain in interior Alaska were studied over a 3-yr period using minirhizotrons placed inside and outside large permanent exclosures. Fine root production and mortality varied seasonally, with greatest rates of production occurring during June each year, and greatest rates of mortality occurring in fall and over winter. Annual production and mortality during 1993, a year of unusually low precipitation, were significantly higher than during either 1992 or 1994. Aboveground herbivory significantly reduced monthly rates of fine root production, and on an annual basis, fine root production of browsed plots (311.4 ± 31.7 mm·tube−1·yr−1) was significantly less than that of unbrowsed plots (453.8 ± 49.8 mm·tube−1·yr−1) when averaged over 3 yr. Because herbivory had less of an effect on monthly or annual rates of fine root mortality, fine root ...

Applications of minirhizotrons to understand root function in forests and other natural ecosystems

Minirhizotrons have proved useful to understand the dynamics and function of fine roots. However, they have been used comparatively infrequently in forests and other natural plant communities. Several factors have contributed to this situation, including anomalous root distributions along the minirhizotron surface and the difficulty of extracting data from minirhizotron images. Technical and methodological advances have ameliorated some of these difficulties, and minirhizotrons have considerable potential to address some questions of long standing interest. These questions include more fully understanding the role of roots in carbon and nutrient cycling, rates of root decomposition, responses to resource availability and the functional significance of interactions between plant roots and soil organisms. Maximizing the potential for minirhizotrons to help us better understand the functional importance of fine roots in natural plant communities depends upon using them to answer only those questions appropriate to both their inherent strengths and limitations.

The relationship between fine root demography and the soil environment in northern hardwood forests

We used minirhizotrons to measure growing-season fine root dynamics at 0–30 and 50–100 cm depths in two northern hardwoods forests. Concomitant measurements of several soil and site environmental v...

Spatial variation in tree root distribution and growth associated with minirhizotrons

Four minirhizotrons were installed in each of three replicate plots in a deciduous forest dominated by Acer saccharum Marsh. The length growth of tree roots along the surface of the minirhizotrons was measured for a period of one year, and the resulting data were analyzed in nested, averaged and pooled arrangements. The analyses of nested data showed that spatial variation in root growth and abundance among minirhizotrons within plots was greater than variation among plots. Averaging data from minirhizotrons within plots prior to analysis reduced variation about plot means, but extensive intraplot variation invalidates this approach on statistical grounds. Both nested and averaged data failed to account for the contribution of individual roots to the mean, and root production rates were consequently overestimated. Pooling the data from the four minirhizotrons reduced variation about the means, and resulted in a more representative estimate of root production rates. The analysis of composited data can be used to incorporate small-scale variation into a single replicate sample in those circumstances where the activity of the root systems of plant communities is the object of study.

Lifespans of fungal rhizomorphs under nitrogen fertilization in a pinyon-juniper woodland

The lifespan of individual microbes in the soil influences nutrient cycling rates as well as population dynamics, but their responses to global change factors such as anthropogenic nitrogen deposition have been challenging to quantify in situ. We used minirhizotron images to track the abundance and turnover rate of individual fungal rhizomorphs under nitrogen fertilization in a pinyon-juniper woodland in New Mexico. We hypothesized that increases in nitrogen availability would alter rhizomorph lifespan and abundance. Sequential images were collected over eight sampling dates from November 1997 to August 1999, and a total of 278 rhizomorphs were examined. We found that neither standing stocks nor lifespans of rhizomorphs differed significantly between treatments. Lifespans of rhizomorphs lasted eleven months on average, indicating that nutrient immobilization in these structures could last for longer than a growing season in these sites.