Determination of sensory bristles and pattern formation in Drosophila: I. A model

A model is presented to explain the formation of the pattern of sensory bristles in Drosophila. The model is based on the idea that precision and reproducibility in pattern formation can be achieved by averaging out of many moderately accurate responses to positional cues. According to this model, the determination of bristles in imaginal discs occurs in two steps. First, large numbers of imaginal cells synthesize a freely diffusible inducer, the chaetogen. Second, cells in which the concentration of this chaetogen reaches a threshold are induced to differentiate into a bristle apparatus. Induced cells prevent neighboring cells from being induced too. The synthesis of chaetogen is supposed to be a probabilistic response of cells to positional cues, so that a cell located in one region of the disc is more likely to have its chaetogen gene turned on than a cell located in another region. Various probability distributions are shown to generate the various bristle patterns observed in the adult: precisely located bristles (e.g., thoracic macrochaetes), evenly spaced bristles (e.g., tergal microchaetes), and rows of bristles (e.g., thoracic microchaetes). In the particular case of the precisely located bristles, we show that (i) the distribution of chaetogen concentration in the tissue presents a unique maximum even when a large number of contiguous cells are all engaged in the synthesis of chaetogen; (ii) the position of the maximum is largely unaffected by statistical fluctuations in the decision of each cell to synthesize or not to synthesize the chaetogen; (iii) different maxima can be reproducibly generated even when the corresponding populations of chaetogen-producing cells overlap.     

An approximate stochastic optimal control framework to simulate nonlinear neuro-musculoskeletal models in the presence of noise

Optimal control simulations have shown that both musculoskeletal dynamics and physiological noise are important determinants of movement. However, due to the limited efficiency of available computational tools, deterministic simulations of movement focus on accurately modelling the musculoskeletal system while neglecting physiological noise, and stochastic simulations account for noise while simplifying the dynamics. We took advantage of recent approaches where stochastic optimal control problems are approximated using deterministic optimal control problems, which can be solved efficiently using direct collocation. We were thus able to extend predictions of stochastic optimal control as a theory of motor coordination to include muscle coordination and movement patterns emerging from non-linear musculoskeletal dynamics. In stochastic optimal control simulations of human standing balance, we demonstrated that the inclusion of muscle dynamics can predict muscle co-contraction as minimal effort strategy that complements sensorimotor feedback control in the presence of sensory noise. In simulations of reaching, we demonstrated that nonlinear multi-segment musculoskeletal dynamics enables complex perturbed and unperturbed reach trajectories under a variety of task conditions to be predicted. In both behaviors, we demonstrated how interactions between task constraint, sensory noise, and the intrinsic properties of muscle influence optimal muscle coordination patterns, including muscle co-contraction, and the resulting movement trajectories. Our approach enables a true minimum effort solution to be identified as task constraints, such as movement accuracy, can be explicitly imposed, rather than being approximated using penalty terms in the cost function. Our approximate stochastic optimal control framework predicts complex features, not captured by previous simulation approaches, providing a generalizable and valuable tool to study how musculoskeletal dynamics and physiological noise may alter neural control of movement in both healthy and pathological movements.     

Stability of particle motions in a narrow channel flow

Single rows or two rows of identical circular cylinders spaced regularly in a narrow channel flow have been shown to be capable of steady flow provided the cylinders are located at equal lateral positions and with equal spacings in the flow direction. The stability of such steady flows of circular cylinders is studied for periodic perturbations of the particle positions, assuming that every other cylinder is equally perturbed in lateral position and spacing along the channel. This results in two rows which are not symmetrically placed. The suspending fluid is assumed to be an incompressible Newtonian fluid. It is assumed that no external forces or moments act on the cylinders and the effects of inertia forces on the motion of the fluid and the cylinders are negligible. The velocity field of the suspending fluid and the instantaneous velocities of the cylinders are computed by the finite element method. The translational velocities of the cylinders are obtained for a large number of particle positions, from which the trajectories of their relative motion are determined for various initial positions near the regular single-file and two-file arrangements. It is shown that when the initial arrangements of the cylinders are slightly perturbed from the regular (alternating) two-file flows, the trajectories of their relative motions become small closed loops around the regular two-file arrangements. In contrast, such small closed trajectories are not obtained when they start from the arrangements near the regular single-file flows or regular (symmetric) double-file flows, suggesting that these flows are unstable under the conditions examined.     

Intermittent control models of human standing: similarities and differences

Two architectures of intermittent control are compared and contrasted in the context of the single inverted pendulum model often used for describing standing in humans. The architectures are similar insofar as they use periods of open-loop control punctuated by switching events when crossing a switching surface to keep the system state trajectories close to trajectories leading to equilibrium. The architectures differ in two significant ways. Firstly, in one case, the open-loop control trajectory is generated by a system-matched hold, and in the other case, the open-loop control signal is zero. Secondly, prediction is used in one case but not the other. The former difference is examined in this paper. The zero control alternative leads to periodic oscillations associated with limit cycles; whereas the system-matched control alternative gives trajectories (including homoclinic orbits) which contain the equilibrium point and do not have oscillatory behaviour. Despite this difference in behaviour, it is further shown that behaviour can appear similar when either the system is perturbed by additive noise or the system-matched trajectory generation is perturbed. The purpose of the research is to come to a common approach for understanding the theoretical properties of the two alternatives with the twin aims of choosing which provides the best explanation of current experimental data (which may not, by itself, distinguish beween the two alternatives) and suggesting future experiments to distinguish beween the two alternatives.     

Dopaminergic modulation of the P50 auditory-evoked potential in a computer model of the CA3 region of the hippocampus: its relationship to sensory gating in schizophrenia

 We modeled the neuronal circuits that may underlie a sensory-processing deficit associated with schizophrenia. Schizophrenic patients have small P50 auditory-evoked responses to click stimuli compared to normal subjects. The P50 auditory-evoked response is a positive waveform recorded in the EEG approximately 50 ms after the auditory click stimulus. In addition to relatively small amplitudes, schizophrenic patients do not gate or suppress the P50 auditory-evoked response to the second of two paired-click stimuli spaced 0.5 s apart. Neuropleptic medication, which decreases dopaminergic neuronal transmission, increases the amplitude of the P50 auditory-evoked response but does not improve gating. Normal subjects have large P50 auditory-evoked responses to click stimuli when compared to unmedicated schizophrenic patients, and they gate their response to paired click stimuli or have smaller P50 auditory-evoked response amplitudes to the second of two click stimuli spaced 0.5 s apart. Schizophrenic patients do not gate and have similar response amplitudes to both clicks. We hypothesized that the small amplitudes of unmedicated schizophrenic subjects were due to a state of occlusion whereby excessive background noise in local circuits reduced the ability of cells to respond synchronously to sensory input, thereby reducing the amplitude of the P50 waveform in the EEG. Because the P50 auditory-evoked potential amplitudes increased with neuroleptic medication, which reduces dopaminergic neuronal transmission, we hypothesized a role for dopamine in modulating the signal-to-noise (S/N) in the local circuits responsible for sensory gating. To test the hypothesis that modulation of the S/N ratio reduces sensory gating, we developed a model of the effects of dopaminergic neuronal transmission that modulates the S/N in neuronal circuits. The model uses the biologically relevant computer model of the CA3 region of the hippocampus developed in the companion paper [Moxon et al. (2003) Biol Cybern, this volume]. Modified Hebb cell assemblies represented the response of the network to the click stimulus. The results of our model showed that excessive dopaminergic input impaired the ability of cells to respond synchronously to sensory input, which reduced the amplitudes of the P50 evoked responses. Received: 3 December 2001 / Accepted: 23 October 2002 / Published online: 28 February 2003 Correspondence to: K.A. Moxon (e-mail: karen.moxon@drexel.edu, Tel.: +1-215-8951959, Fax: +1-215-8954983) Supported by USPHS, MH01245 & MH58414, MH-01121, and research grants from the Department of Veterans Affairs and the National Alliance for Research on Schizophrenia and Depression.     

Gradual appearance of a regulated retinotectal projection pattern in Xenopus laevis

The topographic projection pattern formed by the retinal ganglion cell axons in the tectum of the lower vertebrate appears to require positional cues that guide the optic nerve fibers to their appropriate targets. One approach to understanding these positional cues or "positional information" has been to investigate changes in the pattern of the retinotectal projection after surgical manipulation of the embryonic eyebud. Analysis of these apparent changes in the patterns of positional information in the eye, termed "pattern regulation," may provide clues to both the nature of positional information and the mechanisms by which it is assigned to cells in the eyebud. Here we examine pattern regulation in the Xenopus visual system following the replacement of the temporal half of a right eyebud with the temporal half of a left eyebud. This manipulation requires that the left half-eyebud be inverted along its dorsoventral axis. Electrophysiological maps of these compound eyes in postmetamorphic frogs reveal regulated maps; the cells in the temporal half of the NrTl eye project to the tectum with a dorsoventral polarity appropriate for their position in the host eye and not appropriate for the original positions of the grafted cells in the donor eyebud. Paradoxically, the regulated patterns are not apparent in the projections of the original grafted eyebud cells during early larval development. Using fiber-tracing and electrophysiological mapping techniques, we now show that the regulated patterns appear gradually in the projections made by peripheral retinal cells added during mid-larval development. Because the regulation occurs relatively late in development and probably only in the peripheral retinal cells, simple models of epimorphic or morphallactic regulation do not appear to fit this system. Thus, new or more complex models must be invoked to explain the phenomenon of pattern regulation in the developing visual system of Xenopus.     

Perceptual learning reduces interneuronal correlations in macaque visual cortex

Responses of neurons in early visual cortex change little with training and appear insufficient to account for perceptual learning. Behavioral performance, however, relies on population activity, and the accuracy of a population code is constrained by correlated noise among neurons. We tested whether training changes interneuronal correlations in the dorsal medial superior temporal area, which is involved in multisensory heading perception. Pairs of single units were recorded simultaneously in two groups of subjects: animals trained extensively in a heading discrimination task, and "naive" animals that performed a passive fixation task. Correlated noise was significantly weaker in trained versus naive animals, which might be expected to improve coding efficiency. However, we show that the observed uniform reduction in noise correlations leads to little change in population coding efficiency when all neurons are decoded. Thus, global changes in correlated noise among sensory neurons may be insufficient to account for perceptual learning.     

Sexual selection,sexual isolation and the evolution of song in the Drosophila repleta group of species

The evolution of male courtship song pattern in fruit flies of the repleta group within the genus Drosophila is described. We suggest that the archetypal song was composed of two distinct components, an ‘A’ song with short, regular pulse trains which occurred at the beginning of courtship, and a ‘B’ song consisting of longer pulse trains of more widely spaced pulses occurring later in courtship. During evolution of the repleta group some species have lost the A song, others the B song and, in many species, the latter has become less regular and more complex in form. A songs appear to be involved primarily in species recognition and have been subject to stabilizing selection while the B songs sexually stimulate females and have therefore evolved rapidly through the action of sexual selection. Other factors influencing the evolution of song patterns have been certain physiological and energetic constraints.     

The role of microtubules in establishing nuclear spatial patterns in multinucleate green algae

Summary In uninucleate cells, cytokinesis follows karyokinesis, thereby reestablishing a specific nucleus-to-cytoplasm ratio. In multinucleate cells, cytokinesis is absent or infrequent; no plasmalemma boundary defines the cytoplasmic territory of an individual nucleus. Several genera of large multinucleate green algae were examined with epifluorescence light microscopy to determine whether the patterns of cytoplasmic organization establish nuclear cytoplasmic domains. Randomly spaced nuclei, singular mitotic events and cytoplasmic streaming characterize the organization of two genera,Derbesia andBryopsis (Caulerpales). The cells ofValonia, Valoniopsis, Boergesenia, Ventricaria (Siphonocladales), andHydrodictyon (Chlorococcales) display regularly spaced nuclei which undergo synchronous divisions in a stationary cytoplasm. In the cytoplasm of genera with regularly spaced nuclei, microtubules radiate from all nuclei in late telophase-early interphase. These internuclear microtubule arrays are not found in algal genera with randomly spaced nuclei. It is hypothesized that these microtubule arrays play a role in establishing the cytoplasmic domain of each nucleus in genera with regularly spaced nuclei. Loss of microtubule arrays during the events of mitosis correlated positively with the increasing randomization of nuclear patterns in algae grown in microtubule inhibitors. Cytoplasmic domains were maintained when cells were grown in the same media in the dark. This suggests that, after a round of division, regular nuclear spacing in certain multinucleate algae is reestablished by internuclear microtubule arrays, which are not, however, required to maintain spacing during interphase.Dedicated to the memory of Professor Oswald Kiermayer     

An extremal criterion for epimorphic regeneration

Many developing systems obey the principle of continuity: a morphogenetic field, when perturbed, tends to restore the normal local pattern of structures in its organ district. We have investigated physical field theories for a morphogenetic field, seeking constraints which would make a field theory produce the principle of continuity. We assume that during embryonic (ontogenetic) development a leg develops a pattern of positional values and a length which extremize a time-independent functional—the integral, over the length of the leg, of a function of positional values and position. For a single state variable which represents positional value, if a unique extremizing solution for the ontogenetically generated pattern and the length exists, and if no position-dependent functions other than the state variable appear in the integrand, then the principle of continuity is valid: in any regenerated leg the state variable is continuous and each region is locally identical to a region of the ontogenetically generated leg. This proposition is applied to three simple examples. For an exponential gradient and a Jacobi elliptic function there is a set of parameter values and boundary values for which a functional is minimized and the ontogenetically generated leg has an optimal length. Thus a leg which meets these constraints will obey the principle of continuity. However, a functional which when extremized gives a sinusoidal pattern does not in general provide a unique extremal length. Mathematical conditions are discussed under which an ontogenetically generated limb or a regenerated limb represents an asymptotically stable steady state. For a specific model of the transient dynamics in the exponential gradient case, the steady state gradient is asymptotically stable.     

Analysis of periodic patterns in amino acid sequences: Collagen

Methods are given for analyzing regularly spaced patterns of amino acids in proteins and applied to the α₁ chain of collagen. Fourier methods use the transform of the sequence either embedded in a very long array or folded onto a fundamental base period. Filtering through a moveable “window” of definite width is used to display almost regular features at any chosen frequency. A pattern detection method is described for patterns of general shape. Collagen has statistically significant periodicities at fractions of the stagger distance D = 670 Å. Hydrophobic groups show strong orders of 5, 6, 11; proline 5; charged groups 6, 18, 21. Charged residues mostly occur as neutral pairs. Their distribution has strong 6th and 21st orders which also appear in the changes which are paired at multiples of D. Charge pairs separated by (D + 3) residues show a strong 5D/89 pattern and may form a system of salt bridges across the fibril. There is no sign of any regular pattern of amino acids over the triple helix with a period close to its natural pitch of 30 residues. Supercoiled models with six relative turns of the contact edge between paired triple-helical strands are examined.     

Fluctuations in Neuronal Synchronization in Brain Activity Correlate with the Subjective Experience of Visual Recognition

The scientific study of subjective experience is a current major research area in the neurosciences. Coordination patterns of brain activity are being studied to address the question of how brain function relates to behaviour, and particularly methods to estimate neuronal synchronization can unravel the spatio-temporal dynamics of the transient formation of neuronal assemblies. We report here a biophysical correlate of subjective experience. Subjects visualised figures with different levels of noise, while their brain activity was recorded using magnetoencephalography (MEG), and reported the moment in time (corresponding to a noise level) of figure recognition, which varied between individuals, as well as the moment when they saw the figure more clearly, which was mostly common among the participants (thus less subjective). This latter moment is considered to represent psychophysical stochastic resonance (PSR). Fluctuations in neuronal synchronization, quantified using a diffusion coefficient, were lower in occipital cortex when subjects recognised the figure, for a certain noise level, but did not correlate with the moment of PSR. A different pattern was observed in frontal cortex, where lower values of the diffusion coefficient in neuronal synchronization was maintained from the moment of recognition to the moment of PSR. No specific pattern was found analysing signals from temporal or parietal cortical areas. These observations provide support for distinct synchronization patterns in different cortical areas, and represent another demonstration that the subjective, first-person perspective is accessible to scientific methods.      

Projection pattern of sensory neurons in the central nervous system of a homeotic mutation of the moth Manduca sexta

Octopod (Octo) is a mutation of the moth Manduca sexta, which transforms the first abdominal segment (A1) in the anterior direction. Mutant animals are characterized by the appearance of homeotic thoracic-like legs on A1. We exploited this mutation to determine what rules might be used in specifying the fates of sensory neurons located on the body surface of larval Manduca. Mechanical stimulation of homeotic leg sensilla did not cause reflexive movements of the homeotic legs, but elicited responses similar to those observed following stimulation of ventral A1 body wall hairs. Intracellular recordings demonstrated that several of the motoneurons in the A1 ganglion received inputs from the homeotic sensory hairs. The responses of these motoneurons to stimulation of homeotic sensilla resembled their responses to stimulation of ventral body wall sensilla. Cobalt fills revealed that the mutation transformed the segmental projection pattern of only the sensory neurons located on the ventral surface of A1, resulting in a greater number with intersegmental projection patterns typical of sensory neurons found on the thoracic body wall. Many of the sensory neurons on the homeotic legs had intersegmental projection patterns typical of abdominal sensory neurons: an anteriorly directed projection terminating in the third thoracic ganglion (T3). Once this projection reached T3, however, it mimicked the projections of the thoracic leg sensory neurons. These results demonstrate that the same rules are not used in the establishment of the intersegmental and leg-specific projection patterns. Segmental identity influences the intersegmental projection pattern of the sensory neurons of Manduca, whereas the leg-specific projections are consistent with a role for positional information in determining their pattern. © 1995 John Wiley & Sons, Inc.     

Predicting the Responses of Repetitively Firing Neurons to Current Noise

We used phase resetting methods to predict firing patterns of rat subthalamic nucleus (STN) neurons when their rhythmic firing was densely perturbed by noise. We applied sequences of contiguous brief (0.5–2 ms) current pulses with amplitudes drawn from a Gaussian distribution (10–100 pA standard deviation) to autonomously firing STN neurons in slices. Current noise sequences increased the variability of spike times with little or no effect on the average firing rate. We measured the infinitesimal phase resetting curve (PRC) for each neuron using a noise-based method. A phase model consisting of only a firing rate and PRC was very accurate at predicting spike timing, accounting for more than 80% of spike time variance and reliably reproducing the spike-to-spike pattern of irregular firing. An approximation for the evolution of phase was used to predict the effect of firing rate and noise parameters on spike timing variability. It quantitatively predicted changes in variability of interspike intervals with variation in noise amplitude, pulse duration and firing rate over the normal range of STN spontaneous rates. When constant current was used to drive the cells to higher rates, the PRC was altered in size and shape and accurate predictions of the effects of noise relied on incorporating these changes into the prediction. Application of rate-neutral changes in conductance showed that changes in PRC shape arise from conductance changes known to accompany rate increases in STN neurons, rather than the rate increases themselves. Our results show that firing patterns of densely perturbed oscillators cannot readily be distinguished from those of neurons randomly excited to fire from the rest state. The spike timing of repetitively firing neurons may be quantitatively predicted from the input and their PRCs, even when they are so densely perturbed that they no longer fire rhythmically.     

Distance Estimation Is Influenced by Encoding Conditions

Background

It is well established that foveating a behaviorally relevant part of the visual field improves localization performance as compared to the situation where the gaze is directed elsewhere. Reduced localization performance in the peripheral encoding conditions has been attributed to an eccentricity-dependent increase in positional uncertainty. It is not known, however, whether and how the foveal and peripheral encoding conditions can influence spatial interval estimation. In this study we compare observers'' estimates of a distance between two co-planar dots in the condition where they foveate the two sample dots and where they fixate a central dot while viewing the sample dots peripherally.

Methodology/Principal Findings

Observers were required to reproduce, after a short delay, a distance between two sample dots based on a stationary reference dot and a movable mouse pointer. When both sample dots are foveated, we find that the distance estimation error is small but consistently increases with the dots-separation size. In comparison, distance judgment in peripheral encoding condition is significantly overestimated for smaller separations and becomes similar to the performance in foveal trials for distances from 10 to 16 degrees.

Conclusions/Significance

Although we find improved accuracy of distance estimation in the foveal condition, the fact that the difference is related to the reduction of the estimation bias present in the peripheral conditon, challenges the simple account of reducing the eccentricity-dependent positional uncertainty. Contrary to this, we present evidence for an explanation in terms of neuronal populations activated by the two sample dots and their inhibitory interactions under different visual encoding conditions. We support our claims with simulations that take into account receptive fields size differences between the two encoding conditions.     

Oscillations and oscillatory behavior in small neural circuits

In order to determine the dynamical properties of central pattern generators (CPGs), we have examined the lobster stomatogastric ganglion using the tools of nonlinear dynamics. The lobster pyloric and gastric mill central pattern generators can be analyzed at both the cellular and network levels because they are small, i.e., contain only 25 neurons between them and each neuron and synapse are repeatedly identifiable from animal to animal. We discuss how the biophysical properties of each neuron and synapse in the two circuits act cooperatively to generate two different patterns of sequential activity, how these patterns are altered by neuromodulators and perturbed by noise and sensory inputs. Finally, we show how simplified Hindmarsh–Rose models can be made into analog electronic neurons that mimic the lobster neurons and in addition be incorporated into artificial CPGs with robotic applications.     

Non-linear stimulus-response behavior of the human stance control system is predicted by optimization of a system with sensory and motor noise

We developed a theory of human stance control that predicted (1) how subjects re-weight their utilization of proprioceptive and graviceptive orientation information in experiments where eyes closed stance was perturbed by surface-tilt stimuli with different amplitudes, (2) the experimentally observed increase in body sway variability (i.e. the “remnant” body sway that could not be attributed to the stimulus) with increasing surface-tilt amplitude, (3) neural controller feedback gains that determine the amount of corrective torque generated in relation to sensory cues signaling body orientation, and (4) the magnitude and structure of spontaneous body sway. Responses to surface-tilt perturbations with different amplitudes were interpreted using a feedback control model to determine control parameters and changes in these parameters with stimulus amplitude. Different combinations of internal sensory and/or motor noise sources were added to the model to identify the properties of noise sources that were able to account for the experimental remnant sway characteristics. Various behavioral criteria were investigated to determine if optimization of these criteria could predict the identified model parameters and amplitude-dependent parameter changes. Robust findings were that remnant sway characteristics were best predicted by models that included both sensory and motor noise, the graviceptive noise magnitude was about ten times larger than the proprioceptive noise, and noise sources with signal-dependent properties provided better explanations of remnant sway. Overall results indicate that humans dynamically weight sensory system contributions to stance control and tune their corrective responses to minimize the energetic effects of sensory noise and external stimuli.     

Spatio-Temporal Dynamics of Impulse Responses to Figure Motion in Optic Flow Neurons

White noise techniques have been used widely to investigate sensory systems in both vertebrates and invertebrates. White noise stimuli are powerful in their ability to rapidly generate data that help the experimenter decipher the spatio-temporal dynamics of neural and behavioral responses. One type of white noise stimuli, maximal length shift register sequences (m-sequences), have recently become particularly popular for extracting response kernels in insect motion vision. We here use such m-sequences to extract the impulse responses to figure motion in hoverfly lobula plate tangential cells (LPTCs). Figure motion is behaviorally important and many visually guided animals orient towards salient features in the surround. We show that LPTCs respond robustly to figure motion in the receptive field. The impulse response is scaled down in amplitude when the figure size is reduced, but its time course remains unaltered. However, a low contrast stimulus generates a slower response with a significantly longer time-to-peak and half-width. Impulse responses in females have a slower time-to-peak than males, but are otherwise similar. Finally we show that the shapes of the impulse response to a figure and a widefield stimulus are very similar, suggesting that the figure response could be coded by the same input as the widefield response.     

The TORMOZ gene encodes a nucleolar protein required for regulated division planes and embryo development in Arabidopsis

Embryogenesis in Arabidopsis thaliana is marked by a predictable sequence of oriented cell divisions, which precede cell fate determination. We show that mutation of the TORMOZ (TOZ) gene yields embryos with aberrant cell division planes and arrested embryos that appear not to have established normal patterning. The defects in toz mutants differ from previously described mutations that affect embryonic cell division patterns. Longitudinal division planes of the proembryo are frequently replaced by transverse divisions and less frequently by oblique divisions, while divisions of the suspensor cells, which divide only transversely, appear generally unaffected. Expression patterns of selected embryo patterning genes are altered in the mutant embryos, implying that the positional cues required for their proper expression are perturbed by the misoriented divisions. The TOZ gene encodes a nucleolar protein containing WD repeats. Putative TOZ orthologs exist in other eukaryotes including Saccharomyces cerevisiae, where the protein is predicted to function in 18S rRNA biogenesis. We find that disruption of the Sp TOZ gene results in cell division defects in Schizosaccharomyces pombe. Previous studies in yeast and animal cells have identified nucleolar proteins that regulate the exit from M phase and cytokinesis, including factors involved in pre-rRNA processing. Our study suggests that in plant cells, nucleolar functions might interact with the processes of regulated cell divisions and influence the selection of longitudinal division planes during embryogenesis.