Can meristematic activity determine variation in leaf size and elongation rate among four Poa species? A kinematic study

We studied inherent variation in final leaf size among four Poa spp. that live at different elevations. The average final length of leaf 7 of the main stem of the smallest species (Poa alpina) was only one-half that of the largest species (Poa trivialis); it was correlated with leaf elongation rate, but not with the duration of leaf elongation. A faster rate of leaf elongation rate was associated with (a) larger size of the zone of cell expansion, and (b) faster rates of cell production (per cell file) in the meristem, which in turn were due to greater numbers of dividing cells, whereas average cell division rates were very similar for all species (except Poa annua). Also we found that the proliferative fraction equaled 1 throughout the meristem in all species. It was remarkable that rates of cell expansion tended to be somewhat higher in the species with slower growing leaves. We discuss the results by comparing the spatial and material viewpoints, which lead to different interpretations of the role of cell division. Although the presented data do not strictly prove it, they strongly suggest a regulatory role for cell division in determining differences in growth rate among the present four Poa spp.     

Proliferation controls in a linear growth system: theoretical studies of cell division in the cartilage growth plate

The columnar arrangement of dividing cells in the epiphyseal cartilage plates of growing bones provides a model of a linear proliferation system. One factor which determines the rate of cell production, and hence the rate of growth, is the size of the proliferating population. In this one dimensional system this size is equal to the length of the proliferation zone. Two possible mechanisms for a differentiation control that sets a limit to the length of this zone have been tested in computer simulations. While a diffusion gradient control is consistent with cell kinetic measurements a division limit based on an inheritable growth substance is shown to require further development before the model fits experimental data.Cell division in the columns produces linear clones of cells. If the final length of a bone is set by a limit on the number of divisions that the cartilage stem cells can make, then the number of cells per clone is crucial in determining overall bone growth. The parameters that affect linear clone size have been investigated in computer simulations. Clone size depends largely on the relative division rate of stem cells to proliferation zone cells — but the data on stem cell division rates are generally unreliable.The analysis could be applied to other linear proliferating systems.     

Somatic stability of microsatellite loci in Eastern white pine,Pinus strobus L

Variation at nuclear- and chloroplast-encoded microsatellite loci was studied among and within clonally propagated individuals of Eastern white pine. Total DNA was extracted and assayed from gamete-bearing tissue (megagametophytes) located on six different branch positions on each of 12 individual genets. No within-individual variation was observed among 12 loci studied. Estimates of numbers of mitotic cell divisions required to produce the tissue used as the source of genomic DNA were obtained by combining tree growth and anatomical data. This allowed for the calculation of upper bound estimates of numbers of mutations per locus per somatic cell division. The estimated somatic mutation rate was found to be substantially lower than those published for genomic microsatellite mutation rates in other plant species.     

ADAPTATION, CONSTRAINT, AND COMPROMISE IN AVIAN POSTNATAL DEVELOPMENT

1. This paper discusses factors that influence the evolution of growth rate and determine its variation among species of birds. Growth rate is related to evolutionary fitness through the use of time, energy, and nutrients. In addition, balances between factors favouring rapid growth and those favouring slow growth may be investigated directly by experiment and by comparative observation. 2. David Lack (1968) proposed that the growth rate of the young is the optimum balance between selection for rapid growth to reduce the vulnerable period of development and selection for slow growth to reduce the energy requirements of the young. 3. To test Lack's hypothesis, the growth rates of birds, estimated by fitting sigmoid equations to curves relating weight to age, were surveyed widely from the literature. Among all species examined, growth rate was inversely related to adult weight. Among birds of similar size, most variation in growth rate was related to the degree of maturity of the neonate. Altricial chicks, which depend upon their parents for food and warmth, grow more rapidly than precocial chicks, which are self-sufficient shortly after hatching. Lack's hypothesis, which predicts a direct relationship between growth rate and mortality rate, was not supported. 4. I propose that the key to understanding variation in growth rate among birds lies in the balance between rate of cell proliferation or cell growth, on one hand, and acquisition of mature function, on the other. This idea is consistent with principles of cellular and developmental biology. It is supported by comparisons of (a) the neonates of different species, (b) the individual over the course of the developmental period, and (c) tissues whose use is acquired at different stages of development, wherein more mature individuals or tissues grow more slowly than those with less developed function. 5. Species of birds that are classified as semi-precocial develop precocially but grow rapidly. Although these seemingly violate the general rule relating growth rate to precocity, a closer inspection of their development reveals that they too support the rule. In the Common Tern, the legs, which are the key organ in precocial development, grow at the expected slow rate. The body as a whole grows rapidly because the growth increment of the legs is small and their growth is completed quickly. 6. Growth rates of precocial birds do not decrease abruptly at hatching. This points more to gradual tissue differentiation than to the pattern of procurement and allocation of energy as the primary control for growth rate. 7. Precocious development is favoured when the chicks are capable of self-feeding or when food supplies are distant from the next site and travelling time between one and the other is long. Precocity of the neonates frees both parents to feed at a distant food source. 8. Some species having diets with low levels of protein or other nutrients may grow slowly in order to match nutrient requirements to their availability in the diet. This pattern is indicated especially among the Procellariiformes, which feed an oily diet to their young, and also among tropical fruit-eating birds. 9. Some tropical, pelagically-feeding sea-birds that rear only one offspring at a time may not be able to procure food sufficient to support rapid chick growth. Alternative explanations for slow growth among these species include difficulty in obtaining essential nutrients and more precocious development of activity than in related species having more rapid growth.     

Developmental and genetic origins of murine long bone length variation

If we wish to understand whether development influences the rate or direction of morphological evolution, we must first understand the developmental bases of morphological variation within species. However, quantitative variation in adult morphology is the product of molecular and cellular processes unfolding from embryonic development through juvenile growth to maturity. The Atchley-Hall model provides a useful framework for dissecting complex morphologies into their component parts as a way of determining which developmental processes contribute to variation in adult form. We have examined differences in postnatal allometry and the patterns of genetic correlation between age-specific traits for ten recombinant inbred strains of mice generated from an intercross of LG/J and SM/J. Long bone length is closely tied to body size, but variation in adult morphology is more closely tied to differences in growth rate between 3 and 5 weeks of age. These analyses show that variation generated during early development is overridden by variation generated later in life. To more precisely determine the cellular processes generating this variation we then examined the cellular dynamics of long bone growth plates at the time of maximum elongation rate differences in the parent strains. Our analyses revealed that variation in long bone length is the result of faster elongation rates of the LG/J stain. The developmental bases for these differences in growth rate involve the rate of cell division and chondrocyte hypertrophy in the growth plate.     

Minimal effects of latitude on present-day speciation rates in New World birds

The tropics contain far greater numbers of species than temperate regions, suggesting that rates of species formation might differ systematically between tropical and non-tropical areas. We tested this hypothesis by reconstructing the history of speciation in New World (NW) land birds using BAMM, a Bayesian framework for modelling complex evolutionary dynamics on phylogenetic trees. We estimated marginal distributions of present-day speciation rates for each of 2571 species of birds. The present-day rate of speciation varies approximately 30-fold across NW birds, but there is no difference in the rate distributions for tropical and temperate taxa. Using macroevolutionary cohort analysis, we demonstrate that clades with high tropical membership do not produce species more rapidly than temperate clades. For nearly any value of present-day speciation rate, there are far more species in the tropics than the temperate zone. Any effects of latitude on speciation rate are marginal in comparison to the dramatic variation in rates among clades.     

Bone microstructure and developmental plasticity in birds and other dinosaurs

Patterns of bone microstructure have frequently been used to deduce dynamics and processes of growth in extant and fossil tetrapods. Often, the various types of primary bone tissue have been associated with different bone deposition rates and more recently such deductions have extended to patterns observed in dinosaur bone microstructure. These previous studies are challenged by the findings of the current research, which integrates an experimental neontological approach and a paleontological comparison. We use tetracycline labeling and morphometry to study the variability of bone deposition rates in Japanese quail (Coturnix japonica) growing under different experimental conditions. We compare resulting patterns in bone microstructure with those found in fossil birds and other dinosaurs. We found that a single type of primary bone varies significantly in rates of growth in response to environmental conditions. Ranging between 10-50 microm per day, rates of growth overlap with the full range of bone deposition rates that were previously associated with different patterns of bone histology. Bone formation rate was significantly affected by environmental/experimental conditions, skeletal element, and age. In the quail, the experimental conditions did not result in formation of lines of arrested growth (LAGs). Because of the observed variation of bone deposition rates in response to variation in environmental conditions, we conclude that bone deposition rates measured in extant birds cannot simply be extrapolated to their fossil relatives. Additionally, we observe the variable incidence of LAGs and annuli among several dinosaur species, including fossil birds, extant sauropsids, as well as nonmammalian synapsids, and some extant mammals. This suggests that the ancestral condition of the response of bone to environmental conditions was variable. We propose that such developmental plasticity in modern birds may be reduced in association with the shortened developmental time during the later evolution of the ornithurine birds.     

Does climate variability influence the demography of wild primates? Evidence from long‐term life‐history data in seven species

Earth's rapidly changing climate creates a growing need to understand how demographic processes in natural populations are affected by climate variability, particularly among organisms threatened by extinction. Long‐term, large‐scale, and cross‐taxon studies of vital rate variation in relation to climate variability can be particularly valuable because they can reveal environmental drivers that affect multiple species over extensive regions. Few such data exist for animals with slow life histories, particularly in the tropics, where climate variation over large‐scale space is asynchronous. As our closest relatives, nonhuman primates are especially valuable as a resource to understand the roles of climate variability and climate change in human evolutionary history. Here, we provide the first comprehensive investigation of vital rate variation in relation to climate variability among wild primates. We ask whether primates are sensitive to global changes that are universal (e.g., higher temperature, large‐scale climate oscillations) or whether they are more sensitive to global change effects that are local (e.g., more rain in some places), which would complicate predictions of how primates in general will respond to climate change. To address these questions, we use a database of long‐term life‐history data for natural populations of seven primate species that have been studied for 29–52 years to investigate associations between vital rate variation, local climate variability, and global climate oscillations. Associations between vital rates and climate variability varied among species and depended on the time windows considered, highlighting the importance of temporal scale in detection of such effects. We found strong climate signals in the fertility rates of three species. However, survival, which has a greater impact on population growth, was little affected by climate variability. Thus, we found evidence for demographic buffering of life histories, but also evidence of mechanisms by which climate change could affect the fates of wild primates.     

Real-time lineage analysis reveals oriented cell divisions associated with morphogenesis at the shoot apex of Arabidopsis thaliana

Precise knowledge of spatial and temporal patterns of cell division, including number and orientation of divisions, and knowledge of cell expansion, is central to understanding morphogenesis. Our current knowledge of cell division patterns during plant and animal morphogenesis is largely deduced from analysis of clonal shapes and sizes. But such an analysis can reveal only the number, not the orientation or exact rate, of cell divisions. In this study, we have analyzed growth in real time by monitoring individual cell divisions in the shoot apical meristems (SAMs) of Arabidopsis thaliana. The live imaging technique has led to the development of a spatial and temporal map of cell division patterns. We have integrated cell behavior over time to visualize growth. Our analysis reveals temporal variation in mitotic activity and the cell division is coordinated across clonally distinct layers of cells. Temporal variation in mitotic activity is not correlated to the estimated plastochron length and diurnal rhythms. Cell division rates vary across the SAM surface. Cells in the peripheral zone (PZ) divide at a faster rate than in the central zone (CZ). Cell division rates in the CZ are relatively heterogeneous when compared with PZ cells. We have analyzed the cell behavior associated with flower primordium development starting from a stage at which the future flower comprises four cells in the L1 epidermal layer. Primordium development is a sequential process linked to distinct cellular behavior. Oriented cell divisions, in primordial progenitors and in cells located proximal to them, are associated with initial primordial outgrowth. The oriented cell divisions are followed by a rapid burst of cell expansion and cell division, which transforms a flower primordium into a three-dimensional flower bud. Distinct lack of cell expansion is seen in a narrow band of cells, which forms the boundary region between developing flower bud and the SAM. We discuss these results in the context of SAM morphogenesis.     

GROWTH RATES OF BIRDS IN THE HUMID NEW WORLD TROPICS

The growth curves of 40 species of lowland neotropical birds were fitted by logistic equation. The birds were mostly from Panama, Trinidad and Surinam. The growth constants of the fitted equations (asymptote A and growth rate K) were compared within and among species, and with previously published data on temperate species. Growth parameters of tropical passerines are about as variable within species as they are within temperate species. In both cases, variation in A and K between broods is greater than it is within broods. Panamanian birds breed during the dry-wet transition and conditions for growth apparently improve as rainfall increases. Asymptotes of growth curves are higher, and mortality within broods lower, as the breeding season progresses. Asynchronous hatching and the reduction of brood-size by selective starvation of young is a prominent phenomenon during the early part of the breeding season. Several instances are reported, however, of young persisting in nests with inadequate feeding and greatly subnormal weights. Slowed development under conditions of poor nutrition may be adaptive in the tropics if periods of low food availability are short and allow the possibility of recovery from undernourishment. As a group, neotropical lowland passerines (30 species) grow 23% more slowly than a sample of 51 temperate passerines. Variation of growth rates among these tropical species is similar to variation among temperate species, and it is related to adult body-size the length of the nestling period. Young of tropical and temperate species attain similar asymptotes, relative to adult body-weight, by the end of the nestling period. Hypotheses are advanced which might explain the slower growth rate of tropical species, and tested to the extent available data permits. (1) Because brood-size can be changed only by adding or removing whole young, changes in growth rate could provide finer adjustment of the energy requirements of the young to the feeding capacity of the parents. This model predicts different means and variances for growth rate within groups of species with different clutch-sizes, predictions not supported by available data. (2) Growth rate is shown to increase the maximum energy requirement of a nestling only if K exceeds some value determined by the energy requirement of the young, growth rate should vary in proportion to the level of basal maintenance metabolism. In a small sample of tropical species, rates of basal metabolism were 25% lower than in a comparable sample of temperate species. These data therefore support the hypothesis, although the cause of the lower metabolic rate of the tropical nestlings is not known. (3) Daily periods of hypothermia could reduce the energy requirement of the young and at the same time reduce their growth rate; but observations of body temperatures of tropical nestlings are contrary to this hypothesis. (4) The short day-length of tropical climates reduces the time during which young can assimilate energy relative to their energy expenditures. This model predicts that tropical nestlings would have less productive energy available, (consistent with their reduced growth rates), but it also predicts that arctic birds should grow faster than temperate species, which is not confirmed by available data. (5) The low nitrogen content of fruits may cause the slow growth of a few strictly frugivorous species (Oilbird and Bearded Bellbird), but among other tropical species growth rate is not correlated with the estimated proportion of fruit in the diet.     

Bone Cells in Birds Show Exceptional Surface Area,a Characteristic Tracing Back to Saurischian Dinosaurs of the Late Triassic

Background

Dinosaurs are unique among terrestrial tetrapods in their body sizes, which range from less than 3 gm in hummingbirds to 70,000 kg or more in sauropods. Studies of the microstructure of bone tissue have indicated that large dinosaurs, once believed to be slow growing, attained maturity at rates comparable to or greater than those of large mammals. A number of structural criteria in bone tissue have been used to assess differences in rates of osteogenesis in extinct taxa, including counts of lines of arrested growth and the density of vascular canals.

Methodology/Principal Findings

Here, we examine the density of the cytoplasmic surface of bone-producing cells, a feature which may set an upper limit to the rate of osteogenesis. Osteocyte lacunae and canaliculi, the cavities in bone containing osteocytes and their extensions, were measured in thin-sections of primary (woven and parallel fibered) bone in a diversity of tetrapods. The results indicate that bone cell surfaces are more densely organized in the Saurischia (extant birds, extinct Mesozoic Theropoda and Sauropodomorpha) than in other tetrapods, a result of denser branching of the cell extensions. The highest postnatal growth rates among extant tetrapods occur in modern birds, the only surviving saurischians, and the finding of exceptional cytoplasmic surface area of the cells that produce bone in this group suggests a relationship with bone growth rate. In support of this relationship is finding the lowest cell surface density among the saurischians examined in Dinornis, a member of a group of ratites that evolved in New Zealand in isolation from mammalian predators and show other evidence of lowered maturation rates.     

Heterochronic patterns in primate evolution: evidence from endochondral ossification

Heterochrony (evolutionary modifications in developmental timing and/or rates) is widely recognized as an important agent of morphological change. The adaptive significance of heterochronic changes might lie either in the advantages of the derived morphologies (organ size and shape) or the derived growth parameters themselves (rate and duration of growth). We have tested these hypotheses by comparing the growth rate, the duration of growth and the relative length of the adult tibia in Primates in a phylogenetic context. We report an evolutionary decrease in growth rates (paedochronocline) and an increase in the duration of growth (perachronocline), lying in the cline from the last common ancestor of Primates, passing through the last common ancestor of Haplorhini, that of Catarrhini, to the last common ancestor of the Hominidae. However, the variation in the relative length of the adult tibia does not show any phylogenetic pattern. The derived growth parameters in themselves (slower rate, longer duration) would be of adaptive significance and they would have been selected because a prolonged learning period prior to maturity conferred advantage. The proximate (developmental) causation of differences in bone growth rate were also investigated and it was found that cell production rate in the growth plates rather than the chondrocyte size, underlies the variation in bone growth rate.     

The growth rate of the tarsometatarsus bone in birds

Twisting and bending deformities of the long limb bones of growing birds are common. In domestic poultry they are associated with rapid growth. In order to examine the variation in bone growth rates amongst the Class Aves as a step towards learning about the aetiology of growth deformities, we collected data on this and several other variables from a large sample of species. Adult tarsometatarsal length (ATL) was proportional to adult weight (W)^0.36. The exponent did not differ significantly from 1/3. Mean legginess (L), defined as ATL/W^1/3 (i.e. ATL.W^-1/3), was 80 mm.kg^-1/3 but varied according to the habitat occupied; terrestrial and arboreal birds had relatively longer legs than aquatic and aerial ones. ATL growth rate (GR) averaged 20 mm.d^-1 and was not correlated with W. It was highly influenced by pattern of development, being on average about three times greater in altricial compared with precocial birds of the same adult weight. Amongst birds of the same developmental pattern, GR was linearly related to L. Tarsometatarsal growth rate varied from 0.35 to 60 mm.d^-1 in the Class Aves and there was no evidence of intrinsic limits to GR from our results. How rapidly such bone growth rates are achieved remains to be determined.     

Artificial selection sheds light on developmental mechanisms of limb elongation

Species diversity in limb lengths and proportions is thought to have evolved adaptively in the context of locomotor and habitat specialization, but the heritable cellular processes that drove this evolution within species are poorly understood. In this study, we take a novel “micro‐evo‐devo” approach, using artificial selection on relative limb length to amplify phenotypic variation in a population of mice, known as Longshanks, to examine the cellular mechanisms of postnatal limb development that contribute to intraspecific limb length variation. Cross‐sectional growth data indicate that differences in bone length between Longshanks and random‐bred controls are not due to prolonged growth, but to accelerated growth rates. Histomorphometric and cell proliferation assays on proximal tibial growth plates show that Longshanks’ increased limb bone length is associated with an increased number of proliferative chondrocytes. In contrast, we find no differences in other growth plate cellular features known to underlie interspecific differences in limb bone size and shape, such as the rates of chondrocyte proliferation or the size and number of hypertrophic cells in the growth plate. These data suggest that small differences among individuals in the number of proliferating chondrocytes are a potentially important determinant of selectable intraspecific variation in individual limb bone lengths, independent of body size.     

Endemism, diversity, and the threat of tropical moist forest extinctions

Extinction rates have risen to perhaps 10⁴ the background rate. Much of this increase is due to projected influences of habitat loss on regions of the world with tropical moist forest. This ecosystem, home to a disproportionate amount of global biodiversity and a major regulator of regional and global climate, also faces disproportionately severe threats. In this study, we collect diversity and endemism data for tropical forested countries of the world, along with areal and socioeconomic data. While a correlation between overall numbers of species and endemic species per country is expected, we demonstrate that endemism patterns among birds and mammals remain very strongly convergent even after statistically rendering all countries equal in size and overall species richness and after adjusting for spatial autocorrelation. On a per country basis, mammals are generally more threatened than birds in these tropical moist forested countries. Human population growth rates and rising debt among these nations should be viewed as priorities for amelioration by the developed countries. Reserve network extent is not related to numbers of endemic mammals or birds at this large spatial scale.     

Intramembranous ossification of scleral ossicles in Chelydra serpentina

Scleral ossicles are present in many reptiles, including turtles and birds. In both groups the sclerotic ring situated in the eye is composed of a number of imbricating scleral ossicles or plates. Despite this gross morphological similarity, Andrews (1996. An endochondral rather than a dermal origin for scleral ossicles in Cryptodiran turtles. J. Herpetol. 30, 257-260) reported that the scleral ossicles of turtles develop endochondrally unlike those in birds, which develop intramembranously after a complex epithelial-mesenchymal inductive event. This study re-explores one of the species examined by Andrews in order to determine the mode of ossification of scleral ossicles in turtles. A growth series of Chelydra serpentina embryos, including the stages examined by Andrews, were examined by staining separately for cartilage and bone. Results clearly contradict Andrews (1996) and show that the scleral ossicles of Chelydra serpentina develop similarly to those in birds. That is, they develop intramembranously without a cartilage precursor and are likely induced by transient scleral papillae. The sequence of scleral papillae development is broadly similar, but the papillae themselves are not as distinct as those seen in chicken embryos. This study has important consequences for understanding the homology of scleral ossicles among tetrapods.     

Growth rate variation among passerine species in tropical and temperate sites: an antagonistic interaction between parental food provisioning and nest predation risk

Causes of interspecific variation in growth rates within and among geographic regions remain poorly understood. Passerine birds represent an intriguing case because differing theories yield the possibility of an antagonistic interaction between nest predation risk and food delivery rates on evolution of growth rates. We test this possibility among 64 Passerine species studied on three continents, including tropical and north and south temperate latitudes. Growth rates increased strongly with nestling predation rates within, but not between, sites. The importance of nest predation was further emphasized by revealing hidden allometric scaling effects. Nestling predation risk also was associated with reduced total feeding rates and per-nestling feeding rates within each site. Consequently, faster growth rates were associated with decreased per-nestling food delivery rates across species, both within and among regions. These relationships suggest that Passerines can evolve growth strategies in response to predation risk whereby food resources are not the primary limit on growth rate differences among species. In contrast, reaction norms of growth rate relative to brood size suggest that food may limit growth rates within species in temperate, but not tropical, regions. Results here provide new insight into evolution of growth strategies relative to predation risk and food within and among species.     

Rates of Cell Division in the Shoot Apical Meristem of Pisum

The relative rates of cell division in different regions ofthe pea shoot apical meristem were obtained by measuring theincrease in the numbers of metaphases following applicationof colchicine to the plants. Absolute values for the rates ofcell division could be calculated since the average rate ofcell division for the whole apex was known. Measurements ofthe rates of cell division were obtained at defined intervalsduring the course of a single plastochron. Within each regionof the apex the rate of cell division did not change more thanabout two-fold throughout the plastochron. There was very littleor no increase in the rate of cell division associated withleaf initiation. The formation of a leaf primordium and thesubsequent growth of the apical dome apparently result fromchanges in the direction of growth rather than changes in therates of growth. Three main regions were discernible withinthe apical meristem: a region with a slow rate of cell divisionin the apical dome, a region of a faster rate of cell divisionat the base of the apical dome and at the site of initiationof procambial strands, and a region of an intermediate rateof cell division in the newly initiated leaf primordium andthe adjacent part of the shoot axis.     

The relationship between cell size and cell division in the yeast Saccharomyces cerevisiae.

Cell numbers in synchronous cultures of yeast cultured at fast growth rates increase from N to 2N after the first division and from 2N to 4N after the second division. At these fast growth rates, there are equal numbers of parents and daughters. In contrast, at slow growth rates the cell number increases from N to 2N after one division and from 2N to 3N rather than 4N after the second division. Moreover, the percentage of daughters increases with decreasing growth rate. Thus, slowly growing cultures actually consist of two sub-populations having different cell cycle transit times. These observations are predicted if a yeast cell requires a critical size before a particular cell cycle event can be completed and that after completion of this event cell division occurs following a period of time independent of growth rate.     

The decrease in growth of phosphorus-deficient maize leaves is related to a lower cell production

The spatial distribution of leaf elongation and adaxial epidermal cell production in leaf 6 of maize (Zea mays L. cv. Cecilia) plants grown in a growth chamber under two contrasting availabilities of P in the soil was investigated. Lower displacement velocities from 32.5 mm from leaf base and a shorter growth zone were found in low P (LP) leaves compared with control leaves. P deficiency significantly diminished maximum relative elemental growth rate and shifted its location closer to the leaf base. Cells were significantly longer in LP than in control leaves for all positions from the leaf base except at the end of the growth zone. For both treatments it took a similar time for a cell situated at the leaf base to reach the limit of the growth zone. The average length of the cell division zone was decreased by 21% in LP leaves. Significant differences were found in cell production and cell division rates from 12.5 mm from the leaf base although maximum values were similar between P treatments. A shorter zone of cell division with lower cell production rates along most of its length was the regulatory event that decreased cell production, and ultimately leaf elongation rates, in P‐deficient maize plants.