Regeneration of a dominant Mediterranean dwarf-shrub after fire

Abstract. The regeneration after fire of the dominant east Mediterranean dwarf-shrub Sarcopoterium spinosum was studied under various habitat and geographic conditions in northern Israel, northern Crete and the Aegean coast of Turkey. Age structures of regenerating stands were determined 3–9 yr after fire on 13 1 mx 1 m sample stands by analysis of growth rings in the taproot crown of 874 plants. S. spinosum regenerated either by massive seedling recruitment in some habitats or by resprouting from surviving meristems in others. This versatility facilitates adaptation of the species to widely different environmental conditions and is a factor in the maintenance of its dominance after fire at occupied sites. In the years after fire, the growth of younger plants within the stand at all sites was severely suppressed by the older shrubs. Consequently, the age structure of the stands was dominated by one or two older cohorts that became established soon after fire. In addition, on most habitats, older shrubs (usually older than 6 yr) were often composed of clusters of plants that suggested clonal proliferation of ramets. The strongly peaked age structure, together with the appearance of clonal regeneration on the older plants, suggests that subsequent maintenance of the stand was not dependent on recurrent seedling recruitment. This can explain the persistence of S. spinosum in stands of herbaceous vegetation despite the extreme sensitivity of the seedlings to competition under such conditions.     

Effect of waterlogging on regeneration in the dwarf birch (<Emphasis Type="Italic">Betula nana</Emphasis>)

The aim of this study was to determine the effect of waterlogging on annual regeneration in the dwarf birch (Betula nana L.), and to determine which demographic factors are responsible for the absence of B. nana in the waterlogged parts of a peat-bog. Growth parameters and demographic data were collected from 2001 to 2003 at four sites in the Linje peat bog in the Chełmno Lake District in northern Poland. Three of the sites represented waterlogged open bog communities. The fourth site served as the reference. The density of first-year ramets, annual shoot growth, and the proportion of flowering shoots was significantly lower at the waterlogged sites than at the reference site. No seedlings were found during the course of the study, even though flowering and fruit production took place at the reference site. The flowering shoots were mainly between nine and thirteen years old. Ramets in this age range were found only at the reference site due to high mortality in mature ramets at the waterlogged sites. Mean ramet height was significantly lower at the waterlogged sites than at the reference site, and was strongly correlated with mean ramet age. The distribution of B. nana in peat bogs is limited primarily by two demographic factors: the high mortality rate in older ramets, and the low rate of vegetative reproduction. Another contribution factor is that the proportion of older fertile ramets is low. The effects of fluctuations in the water table on distribution of B. nana are also discussed.     

Senescence in growth and reproductive allocation in a bunchgrass

• Senescence is a puzzling phenomenon. Few convincing studies of senescence in perennial herbaceous plants exist. While ramets are known to senesce, whether senescence of bunchgrasses actually occurs is not clear.
• In this study, we grew a set of plants of Elymus excelsus, a bunchgrass, to examine plant size, sexual reproduction and bud formation in individual plants in relation to their gradual ageing, in order to determine whether E. excelsus experiences senescence. We collected data in two consecutive years (2009 and 2010) from field samples of plants from 1 to 5 years old. Using regression models, we performed age‐related analyses of growth and reproduction parameters.
• Our results showed that individual plant size (diameter, individual biomass), total biomass of ramets, number and biomass of reproductive ramets, percentage of ramets that were reproductive, reproductive allocation, over‐wintering buds and juvenile ramets all declined with age. However, vegetative growth (number and biomass of vegetative ramets) did not decrease with age.
• Those plants that survived, dwindled in size as they aged. However, no plants shifted their resource allocation between growth and reproduction as they aged, so the shift in allocation did not account for the fall in size.

     

Variation of carbon age of fine roots in boreal forests determined from 14C measurements

Background and aims

The main objectives of this study were to determine how the carbon age of fine root cellulose varies between stands, tree species, root diameter and soil depth. In addition, we also compared the carbon age of fine roots from soil cores of this study with reported values from the roots of the same diameter classes of ingrowth cores on the same sites.

Methods

We used natural abundance of ¹⁴C to estimate root carbon age in four boreal Norway spruce and Scots pine stands in Finland and Estonia.

Results

Age of fine root carbon was older in 1.5–2 mm diameter fine roots than in fine roots with <0.5 mm diameter, and tended to be older in mineral soil than in organic soil. Fine root carbon was older in the less fertile Finnish spruce stands (11–12 years) than in the more fertile Estonian stand (3 and 8 years), implying that roots may live longer in less fertile soil. We further observed that on one of our sites carbon in live fine roots with the 1.5–2 mm diameter was of similar C age (7–12 years) than in the ingrowth core roots despite the reported root age in the ingrowth cores – being not older than 2 years.

Conclusions

From this result, we conclude that new live roots may in some cases use old carbon reserves for their cellulose formation. Future research should be oriented towards improving our understanding of possible internal redistribution and uptake of C in trees.     

How past and present influence the foraging of clonal plants?

Clonal plants spreading horizontally and forming a network structure of ramets exhibit complex growth patterns to maximize resource uptake from the environment. They respond to spatial heterogeneity by changing their internode length or branching frequency. Ramets definitively root in the soil but stay interconnected for a varying period of time thus allowing an exchange of spatial and temporal information. We quantified the foraging response of clonal plants depending on the local soil quality sampled by the rooting ramet (i.e. the present information) and the resource variability sampled by the older ramets (i.e. the past information). We demonstrated that two related species, Potentilla reptans and P. anserina, responded similarly to the local quality of their environment by decreasing their internode length in response to nutrient-rich soil. Only P. reptans responded to resource variability by decreasing its internode length. In both species, the experience acquired by older ramets influenced the plastic response of new rooted ramets: the internode length between ramets depended not only on the soil quality locally sampled but also on the soil quality previously sampled by older ramets. We quantified the effect of the information perceived at different time and space on the foraging behavior of clonal plants by showing a non-linear response of the ramet rooting in the soil of a given quality. These data suggest that the decision to grow a stolon or to root a ramet at a given distance from the older ramet results from the integration of the past and present information about the richness and the variability of the environment.     

Micro‐climate determines oviposition site selection and abundance in the butterfly Pyrgus armoricanus at its northern range margin

相似文献   

Comparison of phenotypic and genetic clone delineation in quaking aspen, <Emphasis Type="Italic">Populus tremuloides</Emphasis>

Key message

Clonal delineation at nuclear microsatellites and phenotypic traits showed high correspondence and revealed an important role of both sexual and clonal reproduction for stand genetic structure.

Abstract

Quaking aspen (Populus tremuloides Michx.) grows throughout the northern and central portions of North America. Reproduction occurs both sexually via seeds and clonally from root suckers. Clonal delineation using morphological/phenological traits, and more recently, highly variable nuclear microsatellites have shown considerable variation in the size of clonal assemblies, and the relative importance of sexual versus clonal reproduction across the species range. In order to provide reliable estimates of genet size (N/G; ramets per sampled genet) and genotypic diversity (G/N; genets/ramets), and to compare genetic and phenotypic clone delineation, we characterized 181 sampled stems (ramets) at seven nuclear microsatellites, and morphological and phenological traits from six clones (genet size ≥11). Genotypic diversity was moderate (G/N = 0.18) and within the range reported in other studies across North America. Multivariate statistics revealed a high correspondence between genetic and phenotypic clone delineation, both with and without predefined genetic groups (94.2 %, 81.7 %). Moderate average genet size (5.6 ramets per genet) and the occurrence of genetically distinct single-ramet genets surrounded by larger genets suggested intermediate levels of sexual reproduction contributing to the genetic structure of this stand. Significant differences among genets were found for phenological and morphological traits such as bark thickness and leaf shape. However, most clones showed no significant differences in diameter growth which was likely caused by poor drainage in this high clay soil that inhibited the expression of genetic differences in growth.

     

Ecological correlates of seed mass variation in Phoradendron juniperinum,a xylem-tapping mistletoe

Summary We investigated several ecological correlates of seed mass variation in the hemiparasitic, xylemtapping mistletoe, Phoradendron juniperinum. Mean seed mass varied two-fold among plants between the ages of 4 and 14 years old and was positively correlated with parental plant age. Both the standard deviation and the coefficient of variation in mean seed mass decreased with increasing plant age demonstrating that, on average, younger plants produced seed with more variable mass. Nitrogen concentrations (mg nitrogen per gram of seed) of both the seed and fruit (pericarp) were not correlated with mass or the age of the parent plant from which the seed was taken. However, the nitrogen content per seed (mg nitrogen per seed) was positively correlated with the mean seed dry mass and the age of the seed parent, suggesting that the carbon to nitrogen ratio of individual seeds remained relatively constant as seed mass increased and plants grew older. Seed germination ranged between 20% and 86% and was positively correlated with mass and parental plant age. Heavier seeds (seeds from older plants) also had the highest root radicle growth rates. Furthermore, the final root radicle length after 76 d of growth was positively correlated with seed dry mass. When grown on a medium containing an extract prepared from the host plant foliage, all seeds showed lower germination, grew more slowly and had shorter overall root radicles, but had significantly greater development of the haustorial disks (the holdfast which forms the host-parasite junction in Phoradendron) than seeds grown on a control medium. Our results suggest that, on average, seeds of greater mass produced by older plants have a greater total resource pool per propagule (fruit + seed). This resource pool may be important in conferring a greater potential for dispersal (fruit), survival, colonization, and establishment.     

Hidden crown jewels: the role of tree crowns for bryophyte and lichen species richness in sycamore maple wooded pastures

Tree crowns typically cover the vast majority of the surface area of trees, but they are rarely considered in diversity surveys of epiphytic bryophytes and lichens, especially in temperate Europe. Usually only stems are sampled. We assessed the number of bryophyte and lichen species on stems and in crowns of 80 solitary sycamore maple trees (Acer pseudoplatanus) at six sites in wooded pastures in the northern Alps. The total number of species detected per tree ranged from 13 to 60 for bryophytes, from 25 to 67 for lichens, and from 42 to 104 for bryophytes and lichens considered together. At the tree level, 29 % of bryophyte and 61 % of lichen species were recorded only in the crown. Considering all sampled trees together, only 4 % of bryophyte, compared to 34 % of lichen species, were never recorded on the stem. Five out of 10 red-listed bryophyte species and 29 out of 39 red-listed lichen species were more frequent in crowns. The species richness detected per tree was unexpectedly high, whereas the proportion of exclusive crown species was similar to studies from forest trees. For bryophytes, in contrast to lichens, sampling several stems can give a good estimation of the species present at a site. However, frequency estimates may be highly biased for lichens and bryophytes if crowns are not considered. Our study demonstrates that tree crowns need to be considered in research on these taxa, especially in biodiversity surveys and in conservation tasks involving lichens and to a lesser degree also bryophytes.     

The effects of rooting frequency and position of rooted ramets on plasticity and yield in a clonal species: an experimental study with Glechoma hederacea

Clonal plants produce numerous ramets that can be distributed over a considerable area. Resources are translocated between ramets, especially when they occupy microsites of different quality, or places where leaves or roots cannot be deployed. It is common for a proportion of the ramets of clones and clonal fragments to lack roots. We conducted a greenhouse study using clonal fragments of Glechoma hederacea to examine the effects of differences in the number and position of rooted ramets on yield and plasticity of clonal fragments. We hypothesized that (1) mass of roots and root mass ratio would increase as the number of rooted ramets decreased, (2) plasticity in rooted ramets would buffer the clonal fragment against reduction in yield as the number of rooted ramets declined, (3) ramet plasticity in response to the absence of rooting, and the beneficial effects of this plasticity, would be greater when older ramets were rooted. The same yield was achieved in clonal fragments with only one out of four ramets rooted as in clonal fragments with all four of their ramets rooted, regardless of whether rooting was confined to older or younger ramets. Plasticity in biomass allocated to roots was greater in older rooted ramets succeeded by unrooted ramets than in younger rooted ramets preceded by unrooted ramets. Modular plasticity, involving both direct responses to local conditions, and indirect responses to the conditions experienced by connected modules, buffered performance against variation in rooting ability, enabling clonal fragments to maintain their yield and lateral expansion even when a high proportion of their ramets lacked roots.     

Variation in demographic patterns and population structure of raccoons across an urban landscape

Raccoons (Procyon lotor) are considered synanthropic, with high densities reported from urban landscapes. However, little information is available on population density and demography within the urban matrix. To better understand how urban land-use patterns influence raccoon density and demographic patterns, we sampled raccoons at multiple, replicated sites across an urban landscape. Density differed by land-use type (F_2,17 = 4.66, P = 0.027): urbanized sites, = 4.96 ± 2.64 raccoons/km², range = 1.25–10.00 raccoons/km²; urban open sites, = 14.84 ± 6.35 raccoons/km², range = 3.00–29.25 raccoons/km²; rural open sites, = 15.50 ± 4.66 raccoons/km², range = 13.00–20.25. Although we found no clear patterns in sex ratio, reproductive condition, or body condition, we observed differences in age structure among urban open, rural open, and urbanized sites. The most striking difference was the absence of older animals at urbanized sites and relatively low numbers of young individuals at urban open sites. Raccoons were the dominant mesocarnivore in open fragments, but less so in the urban matrix. Spatial variation in density across urban landscapes is likely influenced by site level differences in abundance of anthropogenic resources and differences in habitat quality. Furthermore, the association between changes in land-use and population age structure may have reflected different mortality sources across the landscape. Our results illustrate that wildlife species considered synanthropic may have complex relationships with urban landscapes. © 2012 The Wildlife Society.     

The use of sectioned otoliths to age barramundi (Lates calcarifer) (Bloch, 1790) [Centropomidae]

The relationship between the number of rings present in sagittal otoliths and the age of barramundi, Lates calcarifer (Bloch, 1790) [Centropomidae], was investigated by examining cross sectioned otoliths of 37 tagged fish of known age between 1 and 5 years from the Johnstone River, north Queensland. Concentric rings were clearly visible in all otolith sections and were validated as annual marks. The technique was then used to estimate the age and calculate von Bertalanffy growth parameters for 70 barramundi from the Fitzroy River, central Queensland. Growth appeared to be rapid but variable in the first year; the von Bertalanffy growth parameters for length versus age were L =690 mm, K=0.53, t ₀= 0.003 years. October 1 was designated as the birth date. Whole otolith length, width and thickness were also approximated well by the von Bertalanffy equation. We suggest that examination of otoliths is a useful technique for ageing barramundi but note that further validation of the ageing method is still needed for fish older than six years.     

Ramet dynamics in a centrifugally expanding clonal sedge: a matrix analysis

Carex humilis is a clonal sedge that can form distinct rings of densely aggregated ramets. We hypothesize that rings form because both production of new ramets and ramet dispersal are positively correlated to ramet size. This would lead to an overrepresentation of fast-moving and large ramets with high ramet production at the periphery, whereas slow-moving and small ramets with low ramet production would mainly be found in the interior of rings. We use matrix models to analyse how ramet populations both at the periphery and in the interior develop in the absence of ramet dispersal. We found that the stable size class distributions of ramets predicted by the models were not different from the distributions found in the field. Also, the asymptotic ramet population growth rates (λ₁) were the same. Hence, we conclude that rings would form even in the absence of a link between ramet dispersal and ramet production. Further analysis of the matrix models showed that the ramet population increases at the periphery but decreases in the interior of rings because medium and large ramets produce fewer large ramets in the interior than at the periphery. We also found that the temporal variance in λ₁ and transitions rates during the four study years was much higher at the periphery than in the interior. Our results suggest that rings may form because C. humilis ramets use below-ground resources from a much larger area than the one covered by the shoots. As the clone grows larger, the soil volume available to the ramets in the interior decreases because their access to soil outside the ring is cut-off by the ramets at the periphery. Ramet density in the interior is therefore decreasing.     

Plasticity in fitness and fitness-related traits at ramet and genet levels in a tillering grass Panicum miliaceum under patchy soil nutrients

Seedling and root sprout recruitment after a volcanic deforestation was examined in a Prunus ssiori population. According to genotypes of five microsatellite (SSR) loci, 44 genets were identified among 188 ramets ≥ 5 cm DBH in a 2.3-ha plot in 2000. The genets formed mutually exclusive patches. Because only one repeat change in an SSR locus distinguished five genets from adjacent genets, there is a possibility that the five genets derived from somatic mutations as well as sexual reproduction. Thus, at least 39 genets originated from seedlings, and at least 144 ramets derived from root sprouts. The seedlings were recruited after the volcanic eruptions in 1739 and before 1972 because trees of 5 cm DBH were 28 years of age, estimated from annual rings. As the largest ramet recruited from a root sprout was estimated to be 94 years old, root sprout recruitment began between 1739 and 1906. Thus, the estimated minimum rates of recruitment from seedlings and root sprouts were 0.167 y^–1 and 0.618-2.182 y^–1, respectively. The clonal diversity of the P. ssiori population (Simpson's D = 0.92) was higher than that of other clonal plant populations. The inbreeding coefficient was significantly negative (Wright's F _IS = −0.069). These results suggest that the frequent seedling recruitment in an outbreeding system maintains the clonal diversity in the population. This revised version was published online in June 2006 with corrections to the Cover Date.     

Consequences of fission in the coral <Emphasis Type="Italic">Siderastrea siderea</Emphasis>: growth rates of small colonies and clonal input to population structure

Colony age and size can be poorly related in scleractinian corals if colonies undergo fission to form smaller independent patches of living tissue (i.e., ramets), but the implications of this life-history characteristic are unclear. This study explored the ecological consequences of the potential discrepancy between size and age for a massive scleractinian, first by testing the effect of colony origin on the growth of small colonies, and second by quantifying the contribution of ramets to population structure. Using Siderastrea siderea in St. John (US Virgin Islands) as an experimental system, the analyses demonstrated that the growth of small colonies derived from sexual reproduction was 2.5-fold greater than that of small ramets which were estimated to be ≈100 years older based on the age of the parent colonies from which they split. Although fission can generate discrete colonies, which in the case of the study reef accounted for 42% of all colonies, it may depress colony success and reef accretion through lowered colony growth rates.     

The oldest known clones of Salix herbacea growing in the Northern Apennines,Italy are at least 2000 years old

Premise

Dominant in many ecosystems around the world, clonal plants can reach considerable ages and sizes. Due to their modular growth patterns, individual clonal plants (genets) can consist of many subunits (ramets). Since single ramets do not reflect the actual age of genets, the ratio between genet size (radius) and longitudinal annual growth rate (LAGR) of living ramets is often used to approximate the age of clonal plants. However, information on how the LAGR changes along ramets and how LAGR variability may affect age estimates of genets is still limited.

Methods

We assessed the variability of LAGR based on wood-section position along the ramets and on the duration of the growing season on three genetically distinct genets of Salix herbacea growing in the Northern Apennines (Italy). We compared genet ages estimated by dividing genet radius by the LAGRs of its ramets.

Results

LAGR increased significantly from the stem apex to the root collar; indicating that ramet growth rate decreased with time. Furthermore, a difference of ca. 2 weeks in the onset of the growing period did not impact LAGR. Considering the high LAGR variability, we estimated that the three genets started to grow between ~2100 and ~7000 years ago, which makes them the oldest known clones of S. herbacea even considering the most conservative age estimate.

Conclusions

Our findings indicate that analyzing ramets at the root collar provides an integrative measurement of their overall LAGR, which is crucial for estimating the age of genets.     

Poterium Spinosum L. Elemento Del Mediterraneo Orientale A Calamosca (Capo S. Elia,Sardegna)

Abstract

«Poterium spinosum» L. in Western Mediterranean at Calamosca (Capo S. Elia, Sardegna). — The Author has studied the station of Poterium spinosum at Calamosca, a small valley of S. Elia Cape, near Cagliari, Sardinia. Im Sardinia this place is the only one where P. spinosum grows on the contrary to the information given in RIKLI (1946) where P. spinosum is indicated as growing everywhere along the coast in the island. As it is known, P. spinosum is very common along the coast in Greece, Lebanon, Israel, Syria; is absent in Egypt; is present in the calcareous part of Cyrenaica, in Sicily between Siracusa and Capo Passero; it reaches westwards the small island of Djerba (Tunisia) and Calamosca (S. Elia Cape — Sardinia). In Dalmatia has been reported for Spalato (Split) and Ragusa (Dubrovnik). It has not been found lately at Fiume (Rijeka) neither at Tivoli (Latium), Cotrone (Basilicata) and Bari (Apulia). P. spinosum at Calamosca in the higher part of the station is tipically rock-dwelling, lower is still rock-dwelling but associated with Thymus capitatus H. et Lk., while in the lowert part of the station P. spinosum grows on gravel with Thymelaea hirsuta Endl. P. spinosum at Calamosca has never been found growing with Thymus capitatus and Thymelaea hirsuta both. The center of the distribution area of P. spinosum is in eastern Mediterranean where it forms (Greece and Greek islands) a typical association with Thymelaea hirsuta (RIKLI 1946). In Israel P. spinosum is associated with Thymus capitatus (ZOHARY). The distribution area of P. spinosum is solid in its eastern part, fragmented in central Mediterranean and sparse towards the west. Recently the species, which has 2n = 28 chromosomes (LARSEN 1955), has been found to have hypoglycemic principles active against diabetes.     

Gender Differences in Health Expectancies across the Disablement Process among Older Thais

Objectives

To estimate health expectancies based on measures that more fully cover the stages in the disablement process for the older Thais and examine gender differences in these health expectancies.

Methods

Health expectancies by genders using Sullivan’s method were computed from the fourth Thai National Health Examination Survey conducted in 2009. A total of 9,210 participants aged 60 years and older were included in the analysis. Health measures included chronic diseases; cognitive impairment; depression; disability in instrumental activities of daily living (IADL); and disability in activities of daily living (ADL).

Results

The average number of years lived with and without morbidity and disability as measured by multiple dimensions of health varied and gender differences were not consistent across measures. At age 60, males could expect to live the most years on average free of depression (18.6 years) and ADL disability (18.6 years) and the least years free of chronic diseases (9.1 years). Females, on the contrary, could expect to live the most years free of ADL disability (21.7 years) and the least years free of IADL disability (8.1 years), and they consistently spent more years with all forms of morbidity and disability. Finally, and for both genders, years lived with cognitive impairment, depression and ADL disability were almost constant with increasing age.

Conclusion

This study adds knowledge of gender differences in healthy life expectancy in the older Thai population using a wider spectrum of health which provides useful information to diverse policy audiences.     

Simulations of clonal species genotypic diversity – trembling aspen (Populus tremuloïdes) as a case study

We built two models to follow clonal species genotypic diversity (G/N) over long periods of time at the stand and landscape levels. The models were then validated with empirical data from trembling aspen (Populus tremuloides) populations in Quebec’s boreal forest. Data was collected using a chronosequence approach in seven sites that burned in 1717, 1760, 1797, 1823, 1847, 1944, and 1916. Genetic identification was done by using four microsatellite loci. At the stand scale, simulations were repeated for a genet size of 5, 25, 50 and 100 ramets each. At the landscape level, we simulated the cumulative genet survival rate under different fire cycles (5–500 years) for 500 years after fire. Stand simulations indicated that ramet mortality within genets rather than genet mortality accounts for the increase in G/N with time since fire. Both the initial genet size and the recurrent suckering of some genets (or ramet recruitment) play an important role in maintaining high G/N levels for long periods of time. In general, the larger the number of ramets per genet, the longer the genet survives under a gap disturbance regime and a minimum of 100 ramets per genet is required to maintain aspen genet survival for 500 years. At the landscape level, genet loss increases as the fire cycle gets longer. In Quebec’s boreal forest, short rotation even-aged management practices seem to maintain a genet survival rate similar to that produced by the natural succession regime.     

Changes in metal-binding peptides due to acclimation to cadmium transferred between ramets of Salvinia minima

Summary Different ramets of a clonal plant exploiting a patchily metal-contaminated habitat may be exposed to different levels of toxic metals. This study investigated whether the exposure of older (parent) ramets to Cd affected the levels of metal-binding peptides and essential elements in younger (daughter) ramets which were not exposed to ambient Cd. Pre-treatment of parent ramets of Salvinia minima with 50 g Cd·l^–1 increased the levels of thiols and phytochelatins (PCs), decreased Mg and increased Cu, Zn and S in daughter ramets growing in a Cd-free medium. Acclimation of parents to lower Cd levels (10 and 25 g Cd·l^–1) increased thiols and decreased cysteine and glutathione in daughters, but did not increase PCs. Parental acclimation to all Cd concentrations tested reduced PC production in daughter ramets during subsequent Cd exposure. Daughter ramets from parents pre-treated with 25 g Cd·l^–1 were more Cd tolerant than controls. Although the tolerant daughters contained 35% more thiols than controls, fractionation of tissue Cd showed no difference between the control and tolerant ramets in the fraction of Cd bound by thiol compounds. Thus, while the acclimation of parent plants to Cd increased levels of metal-binding peptides and thiols in daughter ramets, the relationship between these compounds and the Cd tolerance of daughters is unclear.