Cap Meristem Research Articles

A Study of GUS Expression in Arabidopsis as a Tool for the Evaluation of Gene Evolution, Function and the Role of Expression Derived from Gene Duplication.

Gene duplication played a fundamental role in eukaryote evolution and different copies of a given gene can be present in extant species, often with expressions and functions differentiated during evolution. We assume that, when such differentiation occurs in a gene copy, this may be indicated by its maintenance in all the derived species. To verify this hypothesis, we compared the histological expression domains of the three β-glucuronidase genes (AtGUS) present in Arabidopsis thaliana with the GUS evolutionary tree in angiosperms. We found that AtGUS gene expression overlaps in the shoot apex, the floral bud and the root hairs. In the root apex, AtGUS3 expression differs completely from AtGUS1 and AtGUS2, whose transcripts are present in the root cap meristem and columella, in the staminal cell niche, in the epidermis and in the proximal cortex. Conversely, AtGUS3 transcripts are limited to the old border-like cells of calyptra and those found along the protodermal cell line. The GUS evolutionary tree reveals that the two main clusters (named GUS1 and GUS3) originate from a duplication event predating angiosperm radiation. AtGUS3 belongs to the GUS3 cluster, while AtGUS1 and AtGUS2, which originate from a duplication event that occurred in an ancestor of the Brassicaceae family, are found together in the GUS1 cluster. There is another, previously undescribed cluster, called GUS4, originating from a very ancient duplication event. While the copy of GUS4 has been lost in many species, copies of GUS3 and GUS1 have been conserved in all species examined.

Effects of Cadmium Stress on Root and Root Border Cells of Some Vegetable Species with Different Types of Root Meristem.

Cadmium is one of the most toxic heavy metals and can be easily absorbed by plants, affecting root growth. Root border cells (RBCs), that are located in the periphery of the root cap and originate from the root cap meristem, represent a convenient tool to study the toxic effects of Cd on root performance. In this work, vegetables with contrasting types of root apical meristem (RAM) organizations were used. The open RAM organizations included pea and cucumber, and the closed RAM organizations included tomato, chili, and eggplant. The number of RBCs were significantly higher in the species possessing open RAM organization: pea (11,330 cells per root) > cucumber (8200) > tomato (2480) > eggplant (1830) > chili (1320). The same trend was observed for cell viability: pea (61%) > cucumber (59%) > tomato (49%) > eggplant (44%) > chili (42%). Pea and cucumber had higher relative radicle elongation rates and a lower increase in stress-induced accumulation of malondialdehyde (MDA), making them more resistant to Cd stress than the vegetables with close RAM organization. Under Cd treatment, the number and viability of RBCs in vegetables with both types of RAM organization were significantly decreased. However, the decreasing ratio of the number and viability of RBCs in pea and cucumber was higher than in tomato, chili, and eggplant. Taken together, the plants with the open-type RAM are more tolerant to Cd, and it can be speculated that the cadmium tolerance of the vegetables may be correlated with the number and viability of RBCs in response to cadmium stress.

Root Border Cells as a Convenient Single Cell System to Study Plant-Environmental Interactions: A Case Study for Aluminum Tolerance

Root border cells (RBCs) are a group of cells that originated from the root cap meristem, which are developed by genetic regulation and play a variety of biological functions. Being composed of a homologous single cell population with high metabolic activity and intact cell walls, RBCs represent a highly useful tool for studying various aspects of plant mineral absorption and utilization, as well as plant-soil-microbiome interactions in the rhizosphere. Research on RBCs also promise to become a hotspot in the context of understanding root adaptive responses to hostile environments. In order to take advantage of RBCs as an ideal single cell system in plant-environmental interactions, we summarized the production and function of RBCs and built-up the methodology for RBCs culturing, purification, and quantity control for plant research. The latter is done by using a case study of the application of RBCs to study mechanisms of Al toxicity in plants. This work offers plant scientists a new cognition of adopting RBCs as a convenient single cell system for the multidisciplinary research including (but not limited to) plant physiology, development and genetics, nutrition, and stress and adaptation. Root border cells (RBCs) are derived from the root cap and represent a population of living cells with special physiological activity and biological roles that are different from the root cap cells per se. After being separated from the root cap, RBCs become more active in metabolism than the progenitor root cap cells; for example, they incorporate labeled amino acids into protein 2.6-fold more efficiently than the cells of the root cap. In addition, mRNA and protein were differentially expressed between root cap cells and RBCs. Since the production of RBCs is genetically regulated and RBCs played a variety of biological functions in resistance to biotic and abiotic stresses occurred in the rhizosphere, RBCs were suggested as an ideal single cell system for the study the response of plant root cells to nutrient availability, environmental stresses, and in plant-microbial interactions. Some studies revealed that RBCs, which development is regulated by endogenous and exogenous signals, are biologically viable in the majority of higher plant species. This work reviews the research on RBCs in plant environment interaction and describes the case study of RBCs as a convenient single cell system to study plant responses to Al toxicity.

TOPOISOMERASE1α Acts through Two Distinct Mechanisms to Regulate Stele and Columella Stem Cell Maintenance.

TOPOISOMERASE1 (TOP1), which releases DNA torsional stress generated during replication through its DNA relaxation activity, plays vital roles in animal and plant development. In Arabidopsis (Arabidopsis thaliana), TOP1 is encoded by two paralogous genes (TOP1α and TOP1β), of which TOP1α displays specific developmental functions that are critical for the maintenance of shoot and floral stem cells. Here, we show that maintenance of two different populations of root stem cells is also dependent on TOP1α-specific developmental functions, which are exerted through two distinct novel mechanisms. In the proximal root meristem, the DNA relaxation activity of TOP1α is critical to ensure genome integrity and survival of stele stem cells (SSCs). Loss of TOP1α function triggers DNA double-strand breaks in S-phase SSCs and results in their death, which can be partially reversed by the replenishment of SSCs mediated by ETHYLENE RESPONSE FACTOR115 In the quiescent center and root cap meristem, TOP1α is epistatic to RETINOBLASTOMA-RELATED (RBR) in the maintenance of undifferentiated state and the number of columella stem cells (CSCs). Loss of TOP1α function in either wild-type or RBR RNAi plants leads to differentiation of CSCs, whereas overexpression of TOP1α mimics and further enhances the effect of RBR reduction that increases the number of CSCs Taken together, these findings provide important mechanistic insights into understanding stem cell maintenance in plants.

Extracellular DNA is required for root tip resistance to fungal infection.

Plant defense involves a complex array of biochemical interactions, many of which occur in the extracellular environment. The apical 1- to 2-mm root tip housing apical and root cap meristems is resistant to infection by most pathogens, so growth and gravity sensing often proceed normally even when other sites on the root are invaded. The mechanism of this resistance is unknown but appears to involve a mucilaginous matrix or "slime" composed of proteins, polysaccharides, and detached living cells called "border cells." Here, we report that extracellular DNA (exDNA) is a component of root cap slime and that exDNA degradation during inoculation by a fungal pathogen results in loss of root tip resistance to infection. Most root tips (>95%) escape infection even when immersed in inoculum from the root-rotting pathogen Nectria haematococca. By contrast, 100% of inoculated root tips treated with DNase I developed necrosis. Treatment with BAL31, an exonuclease that digests DNA more slowly than DNase I, also resulted in increased root tip infection, but the onset of infection was delayed. Control root tips or fungal spores treated with nuclease alone exhibited normal morphology and growth. Pea (Pisum sativum) root tips incubated with [(32)P]dCTP during a 1-h period when no cell death occurs yielded root cap slime containing (32)P-labeled exDNA. Our results suggest that exDNA is a previously unrecognized component of plant defense, an observation that is in accordance with the recent discovery that exDNA from white blood cells plays a key role in the vertebrate immune response against microbial pathogens.

Inducible expression of Pisum sativum xyloglucan fucosyltransferase in the pea root cap meristem, and effects of antisense mRNA expression on root cap cell wall structural integrity.

Mitosis and cell wall synthesis in the legume root cap meristem can be induced and synchronized by the nondestructive removal of border cells from the cap periphery. Newly synthesized cells can be examined microscopically as they differentiate progressively during cap development, and ultimately detach as a new population of border cells. This system was used to demonstrate that Pisum sativum L. fucosyl transferase (PsFut1) mRNA expression is strongly expressed in root meristematic tissues, and is induced >2-fold during a 5-h period when mitosis in the root cap meristem is increased. Expression of PsFut1 antisense mRNA in pea hairy roots under the control of the CaMV35S promoter, which exhibits meristem localized expression in pea root caps, resulted in a 50–60% reduction in meristem localized endogenous PsFut1 mRNA expression measured using whole mount in situ hybridization. Changes in gross levels of cell wall fucosylated xyloglucan were not detected, but altered surface localization patterns were detected using whole mount immunolocalization with CCRC-M1, an antibody that recognizes fucosylated xyloglucan. Emerging hairy roots expressing antisense PsFut1 mRNA appeared normal macroscopically but scanning electron microscopy of tissues with altered CCRC-M1 localization patterns revealed wrinkled, collapsed cell surfaces. As individual border cells separated from the cap periphery, cell death occurred in correlation with extrusion of cellular contents through breaks in the wall.

Proteins Among the Polysaccharides

Charles Darwin recognized the power of the root cap as a model for plant signalling and behavior, and used it to explore the ways plants sense and respond to diverse stimuli. Over ensuing decades, various groups have reported tantalizing clues regarding the role of a complex extracellular matrix that ensheaths the tip region housing the apical and root cap meristems. In the course of characterizing root tip resistance to infection and injury and the role border cells play in this phenomenon, we confirmed and extended early- and mid-20th century studies reporting enzyme activities secreted from the root cap. Multidimensional protein analysis revealed, in fact, that >100 proteins are actively synthesized and secreted from the root cap and border cells. This 'root cap secretome' appears to be a critical component of root tip resistance to infection. We have developed a microscopic assay to quantify the protein-based extracellular response to dynamic changes in environmental conditions including hydroponic culture, and present the results here. This tool provides a simple, direct measure that can be used to explore the ways border cells may function in the manner of white blood cells to trap, immobilize, and neutralize threats to the growing root tip.

The cyclic nucleotide gated cation channel AtCNGC10 traffics from the ER via Golgi vesicles to the plasma membrane of Arabidopsis root and leaf cells.

BackgroundThe cyclic nucleotide-gated ion channels (CNGCs) maintain cation homeostasis essential for a wide range of physiological processes in plant cells. However, the precise subcellular locations and trafficking of these membrane proteins are poorly understood. This is further complicated by a general deficiency of information about targeting pathways of membrane proteins in plants. To investigate CNGC trafficking and localization, we have measured Atcngc5 and Atcngc10 expression in roots and leaves, analyzed AtCNGC10-GFP fusions transiently expressed in protoplasts, and conducted immunofluorescence labeling of protoplasts and immunoelectron microscopic analysis of high pressure frozen leaves and roots.ResultsAtCNGC10 mRNA and protein levels were 2.5-fold higher in roots than leaves, while AtCNGC5 mRNA and protein levels were nearly equal in these tissues. The AtCNGC10-EGFP fusion was targeted to the plasma membrane in leaf protoplasts, and lightly labeled several intracellular structures. Immunofluorescence microscopy with affinity purified CNGC-specific antisera indicated that AtCNGC5 and AtCNGC10 are present in the plasma membrane of protoplasts. Immunoelectron microscopy demonstrated that AtCNGC10 was associated with the plasma membrane of mesophyll, palisade parenchyma and epidermal cells of leaves, and the meristem, columella and cap cells of roots. AtCNCG10 was also observed in the endoplasmic reticulum and Golgi cisternae and vesicles of 50–150 nm in size. Patch clamp assays of an AtCNGC10-GFP fusion expressed in HEK293 cells measured significant cation currents.ConclusionAtCNGC5 and AtCNGC10 are plasma membrane proteins. We postulate that AtCNGC10 traffics from the endoplasmic reticulum via the Golgi apparatus and associated vesicles to the plasma membrane. The presence of the cation channel, AtCNGC10, in root cap meristem cells, cell plate, and gravity-sensing columella cells, combined with the previously reported antisense phenotypes of decreased gravitropic and cell enlargement responses, suggest roles of AtCNGC10 in modulating cation balance required for root gravitropism, cell division and growth.

Root Border Cell Development is a Temperature-Insensitive and Al-Sensitive Process in Barley

In vivo and in vitro experiments showed that border cell (BC) survival was dependent on root tip mucigel in barley (Hordeum vulgare L. cv. Hang 981). In aeroponic culture, BC development was an induced process in barley, whereas in hydroponic culture, it was a kinetic equilibrium process during which 300-400 BCs were released into water daily. The response of root elongation to temperatures (10-35 degrees C) was very sensitive but temperature changes had no great effect on barley BC development. At 35 degrees C, the root elongation ceased whereas BC production still continued, indicating that the two processes might be regulated independently under high temperature (35 degrees C) stress. Fifty microM Al could inhibit significantly BC development by inhibiting pectin methylesterase activity in the root cap of cv. 2000-2 (Al-sensitive) and cv. Humai 16 (Al-tolerant), but 20 microM Al could not block BC development in cv. Humai 16. BCs and their mucigel of barley had a limited role in the protection of Al-induced inhibition of root elongation, but played a significant role in the prevention of Al from diffusing into the meristems of the root tip and the root cap. Together, these results suggested that BC development was a temperature-insensitive but Al-sensitive process, and that BCs and their mucigel played an important role in the protection of root tip and root cap meristems from Al toxicity.

Lethality of inducible, meristem-localized ectopic β-glucuronidase expression in plants

GUSA fromEscherichia coli, encoded by theuidA gene, has been successfully used as a plant reporter system for more than a decade with no reported deleterious effects. However, when expressed in coordination with a UDP-glucuronosyltransferase isolated from the root cap meristem ofPisum sativum (PsUGT1) at the onset of mitosis, GUSA expression was lethal in pea, alfalfa, andArabidopsis thaliana. These unexpected results indicate that, under some circumstances, using GUSA in plants is incompatible with life and suggest that the cell-specific lethal phenotype might be useful in selecting for genes specifically involved in regulating the G2-M phase of the cell cycle.

The Root Cap: Cell Dynamics, Cell Differentiation and Cap Function

The root cap is a universal feature of angiosperm, gymnosperm, and pteridophyte roots. Besides providing protection against abrasive damage to the root tip, the root cap is also involved in the simultaneous perception of a number of signals – pressure, moisture, gravity, and perhaps others – that modulate growth in the main body of the root. These signals, which originate in the external environment, are transduced by the cap and are then transported from the cap to the root. Root gravitropism is one much studied response to an external signal. In the present paper, consideration is given to the structure of the root cap and, in particular, to how the meristematic initial cells of both the central cap columella and the lateral portion of the cap which surrounds the columella are organized in relation to the production of new cells. The subsequent differentiation and development of these cells is associated with their displacement through the cap and their eventual release, as “border cells”, from the cap periphery. Mutations, particularly in Arabidopsis, are increasingly playing a part in defining not only the pattern of genetic activity within different cells of the cap but also in revealing how the corresponding wild-type proteins relate to the range of functions of the cap. Notable in this respect have been analyses of the early events of root gravitropism. The ability to image auxin and auxin permeases within the cap and elsewhere in the root has also extended our understanding of this growth response. Images of auxin distribution may, in addition, help extend ideas concerning the positional controls of cell division and cell differentiation within the cap. However, firm information relating to these controls is scarce, though there are intriguing suggestions of some kind of physiological link between the border cells surrounding the cap and mitotic activity in the cap meristem. Open questions concern the structure and functional interrelationships between the root and the cap which surmounts it, and also the means by which the cap transduces the environmental signals that are of critical importance for the growth of the individual roots, and collectively for the shaping of the root system.

Root Caps and Rhizosphere

In this paper we discuss recent work on the physiological, molecular, and mechanical mechanisms that underlie the capacity of root caps to modulate the properties of the rhizosphere and thereby foster plant growth and development. The root cap initially defines the rhizosphere by its direction of growth, which in turn occurs in response to gradients in soil conditions and gravity. The ability of the root cap to modulate its environment is largely a result of the release of exudates and border cells, and so provides a potential method to engineer the rhizosphere. Factors affecting the release of border cells from the outer surface of the root cap, and function of these cells and their exudates in the rhizosphere, are considered in detail. Release of border cells into the rhizosphere depends on soil matric potential and mechanical impedance, in addition to a host of other environmental conditions. There is good evidence of unidentified feedback signals between border cells and the root cap meristem, and some potential mechanisms are discussed. Root border cells play a significant mechanical role in decreasing frictional resistance to root penetration, and a conceptual model for this function is discussed. Root and border cell exudates influence specific interactions between plant hosts and soil organisms, including pathogenic fungi. The area of exudates and border cell function in soil is an exciting and developing one that awaits the production of appropriate mutant and transgenic lines for further study in the soil environment.

Developmental and Thermal Regulation of the Maize Heat Shock Protein, HSP101

The plant heat stress protein, Hsp101, and the yeast ortholog, Hsp104, are required to confer thermotolerance in plants and yeast (Saccharomyces cerevisiae), respectively. In addition to its function during stress, Hsp101 is developmentally regulated in plants although its function during development is not known. To determine how the expression of Hsp101 is regulated in cereals, we investigated the Hsp101 expression profile in developing maize (Zea mays). Hsp101 protein was most abundant in the developing tassel, ear, silks, endosperm, and embryo. It was less abundant in the vegetative and floral meristematic regions and was present at only a low level in the anthers and tassel at anthesis, mature pollen, roots, and leaves. As expected, heat treatment resulted in an increase in the level of Hsp101 protein in several organs. In expanding foliar leaves, husk leaves, the tassel at the premeiosis stage of development, or pre-anthesis anthers, however, the heat-mediated increase in protein was not accompanied by an equivalent increase in mRNA. In contrast, the level of Hsp101 transcript increased in the tassel at anthesis following a heat stress without an increase in Hsp101 protein. In other organs such as the vegetative and floral meristematic regions, fully expanded foliar leaves, the young ear, and roots, the heat-induced increase in Hsp101 protein was accompanied by a corresponding increase in Hsp101 transcript level. However, anthers at anthesis, mature pollen, developing endosperm, and embryos largely failed to mount a heat stress response at the level of Hsp101 protein or mRNA, indicating that Hsp101 expression is not heat inducible in these organs. In situ RNA localization analysis revealed that Hsp101 mRNA accumulated in the subaleurone and aleurone of developing kernels and was highest in the root cap meristem and quiescent center of heat-stressed roots. These data suggest an organ-specific control of Hsp101 expression during development and following a heat stress through mechanisms that may include posttranscriptional regulation.

Image Analysis of Maize Root Caps—Estimating Cell Numbers from 2-D Longitudinal Sections

The cap of the primary root of maize produces several thousand border cells that are shed from the outside of the cap each day. Border cell production is important in the penetration of soil by roots, and in influencing the activity of both beneficial and pathogenic organisms in the rhizosphere. To improve understanding of the dynamics of border cell production, it is desirable to know the number of cells in different parts of the root cap. An image analysis procedure was used to quantify cell dimensions and locations in the median longitudinal section of maize (Zea mays L.) root caps. Calculations based on root symmetry were then used to estimate the number of cells in 3-dimensions. Our estimation procedure was tested initially using regular arrays of identical square and hexagonal shapes to represent cells. It was then tested using two different tissues showing analogous arrays: a transverse section through the maize root cap junction, and a transverse section through a barley root. Good linear correlations were obtained between the number of cells estimated and the number of cells actually counted in the microscope. The numbers of cells in the whole maize root cap (8870±390) were then estimated from longitudinal sections. These numbers of cap cells agreed with values that had been estimated for maize by other methods. In the first tier of the cap meristem, ten-times more meristematic cells were located in the cap flanks (>500 cells) than were present in the columella portion. Similarly, only 7% of cells in the outermost layer of the root were associated with the columella region of the cap, a fraction which compared well with previous measurements of sloughed cells extracted from rhizosphere sand. This present technique can be applied to estimate the numbers of cells in any cylindrically symmetrical tissue from two-dimensional sections.

Meristem-specific suppression of mitosis and a global switch in gene expression in the root cap of pea by endogenous signals

Two functionally distinct sets of meristematic cells exist within root tips of pea (Pisum sativum): the root apical meristem, which gives rise to the body of the root; and the root cap meristem, which gives rise to cells that differentiate progressively through the cap and separate ultimately from its periphery as border cells. When a specific number of border cells has accumulated on the root cap periphery, mitosis within the root cap meristem, but not the apical meristem, is suppressed. When border cells are removed by immersion of the root tip in water, a transient induction of mitosis in the root cap meristem can be detected starting within 5 min. A corresponding switch in gene expression throughout the root cap occurs in parallel with the increase in mitosis, and new border cells begin to separate from the root cap periphery within 1 h. The induction of renewed border cell production is inhibited by incubating root tips in extracellular material released from border cells. The results are consistent with the hypothesis that operation of the root cap meristem and consequent turnover of the root cap is self-regulated by a signal from border cells.

Differential Expression of Proteins and mRNAs from Border Cells and Root Tips of Pea.

Many plants release large numbers of metabolically active root border cells into the rhizosphere. We have proposed that border cells, cells produced by the root cap meristem that separate from the rest of the root upon reaching the periphery of the cap, are a singularly differentiated part of the root system that modulates the environment of the plant root by producing specific substances to be released into the rhizosphere. Proteins synthesized in border cells exhibit profiles that are very distinct from those of the root tip (root cap, root meristem, and adjacent cells). In vivo-labeling experiments demonstrate that 13% of the proteins that are abundant in preparations from border cells are undetectable in root tip preparations. Twenty-five percent of the proteins synthesized by border cells in a 1-h period are rapidly excreted into the incubation medium. Quantitative variation in levels of specific marker proteins, including glutamine synthetase, heat-shock protein 70, and isoflavone reductase, also occurs between border cells and cells in the root tip. mRNA differential-display assays demonstrate that these large qualitative and quantitative differences in protein expression are correlated with similarly distinct patterns of gene expression. These observations are consistent with the hypothesis that a major switch in gene expression accompanies differentiation into root border cells, as expected for cells with specialized functions in plant development.

Transmural secretion of a highly-expressed cell surface antigen of oat root cap cells

A monoclonal antibody (MAb), designated 40.1C2.8, has been produced which recognizes an antigen in the outer membrane and cell wall of oat root cap cells. Antibody binding is insensitive to periodate oxidation and sensitive to proteolytic digestion by trypsin, indicating that the epitope is protein, not carbohydrate. Western immunoblots of polypeptide extracts from oat, maize, and bean seedlings revealed that MAb 40.1C2.8 was primarily directed to an epitope on a 40 kDa protein very highly expressed in root cap cells. Root cap cells and mature sloughed cells were heavily stained with the antibody on silver-enhanced immunogold-stained thick sections of intact roots. At the ultrastructural level, the antigen was found on the cell surface of root cap meristem cells. In the root meristem proper, the antigen was only detected in the vacuole, not on the cell surface. As the root cap cells matured, the thickness of the zone containing the antigen increased. In the outer-most peripheral cells, the antigen was found within the inner layer of the wall, but not coextensive with the secreted mucilage which occurs throughout the wall, indicating that there may be a barrier to transmural secretion even in the most mature cells of the root cap.

The Meristem and Quiescent Centre in Cultured Root Apices of the gib-l Mutant of Tomato (Lycopersicon esculentum Mill.)*

Cultured root apices of tomato bearing the gib-1 mutation, which reduces the levels of endogenous gibberellins, grew slower and were thicker than wild-type controls. This was the result of shorter and broader cells in the meristem of the mutant. Cells of both cortex and stele were affected, but this did not cause any alteration to the volume fraction occupied by these two tissues in the root meristem. Root caps were longer in the mutant and there were also more layers of rhizodermis. All these effects could be reproduced in wild-type roots by addition of 0.1 micromolar 2S,3S paclobutrazol (an inhibitor of gibberellin biosynthesis) to the culture medium and could be normalized in mutant roots by 0.1 micromolar GA3. Cell doubling times in the proximal region of the meristem were similar in mutant and wild-type roots, but were faster in both the quiescent centre (QC) and the cap meristem of the mutant. This latter feature of the mutant roots is likely to be the cause of their longer caps, while the faster rate of division in the QC accounts for the additional tiers of cells that were found to build up in the cortical portion of this zone. These additional tiers failed to form in mutant roots grown in GA3, but they could be induced in wild-type roots by 2S,3S paclobutrazol. These results suggest that endogenous gibberellins may be partly responsible for the slow rate of cell growth and proliferation in the QC.

The recovery of the roots of Zea mays L. from various aluminium treatments: Towards elucidating the regulatory processes that underlie root growth control

Controlled environment studies have demonstrated the potential of roots exposed to Al (5 mg/dm 3) for various times (up to 48 hr) to recover and resume primary growth subsequent to their transfer to Al-free recovery solutions. Ultrastructural investigation of the stages involved in recovery showed that the resumption of root elongation rates during recovery coincided with the presence of a morphologically distinctive secretory activity in peripheral root cap cells. This suggested links for this activity in regulating root growth responses. Changes in the volume of the root cap, attributed to reactivation of the cap meristem initially made inactive by Al, and the recovery of root meristem cells involving nucleolar reaggregation and decreased vacuolation were characteristic of the early stages of root recovery, and preceded by several days a resumption of root elongation rates. Recovery was concomitant with root growth stimulation. Activation of the cells of the quiescent centre to form a new cap was not an early response of roots recovering from Al. In contrast, “continuous” root exposure to Al prevented recovery of the primary root meristem and resulted in severe reductions in root elongation rates and a loss of apical dominance. The stages involved in the recovery of roots from Al are used to discuss root growth regulation and the physiology of Al toxicity.

Modelling of the root cap of Zea mays L in relation to temperature

Rates of mitosis in each of the four tiers of the cap meristem of primary roots of Zea mays L. were measured at uniform root length by a stathmokinetic method and compared with rates for other regions of the apices in seedlings growing at between 15 and 35 °C. The highest elemental rate of cap cell production in the proximal tier and in the cap meristem as a whole occurs at 26 °C and the proportion of cells supplied by the proximal tier rises from 48%, at 15 °C to 90% at 35 °C. The number of cells in the whole cap and in its meristem decreases with increasing temperature. This is due to the general reduction in the width of root apices with age and growth in length being faster at the higher temperatures. The slimming of the apex is effected by a decline in the proportion of divisions that are longitudinal to the files of cells at the quiescent centre. This is enhanced by a slight but continuous increase in the rate of mitosis in the quiescent centre the higher the temperature (in contrast with the rest of the root whose maximal rate is at 2S °C). The decline results in an increase in the elemental rate for longitudinal divisions in the cap initials from 0.02 to 0.17 cells per cell per day between 15 and 35 °C and a consequent change in the cell pattern in the cap.