The tiny enslaved genome of a rhizarian alga.
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Biological Evolution↗
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Unique tRNA introns of an enslaved algal cell.
Nucleomorphs are remnant nuclei of eukaryotic, secondary endosymbionts exclusively found in cryptophytes and chlorarachniophytes. The nucleomorph of the cryptophyte Guillardia theta codes for 36 transfer RNA (tRNA) genes, 15 of them predicted to contain introns and 1 pseudo-tRNA. Some of the predicted intervening sequences are manifested at positions not known in Eukarya, even tRNAs with more than one intron were suggested. By isolating reverse transcriptase-polymerase chain reaction products of the spliced tRNAs we verify the processing of all predicted intron-harboring tRNAs and demonstrate the splicing of the smallest introns (3 nt) investigated so far. However, the spliced intervening sequences are in some cases shifted in respect to the predicted ones. Moreover, we show that introns, if inserted into the B-box of tRNA genes in the nucleomorph of cryptophytes, mimic promoter regions and do not abolish transcription by RNA polymerase III. Consequently, internal nucleomorph-encoded tRNA promoter regions are in some cases dissected from the sequence of the mature tRNAs. By reanalyzing tRNA introns of a recently sequenced red algae we furthermore show that splicing of introns at unusual positions may be introduced in cryptophytes by its secondary endosymbiont. However, in contrast to the rest of the symbiont genome, introns are not minimized in quantity but are instead scattered along the tRNA genes.
Base Sequence↗
Fundamental sensitivity propagators in dissipative systems with a statistical enslaving of fast-relaxing variables.
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Journal Article↗
The ever-expanding information colossus: benefactor or enslaver?
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Computers↗
A new kind of trafficking: child beggars in Asia. Scared, hungry, enslaved.
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Adolescent↗
["Panis", a captive enslaved in the Indian Wars].
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History, 18th Century↗
Coordinated force production in multi-finger tasks: finger interaction and neural network modeling.
During maximal voluntary contraction (MVC) with several fingers, the following three phenomena are observed: (1) the total force produced by all the involved fingers is shared among the fingers in a specific manner (sharing); (2) the force produced by a given finger in a multi-finger task is smaller than the force generated by this finger in a single-finger task (force deficit); (3) the fingers that are not required to produce any force by instruction are involuntary activated (enslaving). We studied involuntary force production by individual fingers (enslaving effects, EE) during tasks when (an)other finger(s) of the hand generated maximal voluntary pressing force in isometric conditions. The subjects (n = 10) were instructed to press as hard as possible on the force sensors with one, two, three and four fingers acting in parallel in all possible combinations. The EE were (A) large, the slave fingers always producing a force ranging from 10.9% to 54.7% of the maximal force produced by the finger in the single-finger task; (B) nearly symmetrical; (C) larger for the neighboring fingers; and (D) non-additive. In most cases, the EE from two or three fingers were smaller than the EE from at least one finger (this phenomenon was coined occlusion). The occlusion cannot be explained only by anatomical musculo-tendinous connections. Therefore, neural factors contribute substantially to the EE. A neural network model that accounts for all the three effects has been developed. The model consists of three layers: the input layer that models a central neural drive; the hidden layer modeling transformation of the central drive into an input signal to the muscles serving several fingers simultaneously (multi-digit muscles); and the output layer representing finger force output. The output of the hidden layer is set inversely proportional to the number of fingers involved. In addition, direct connections between the input and output layers represent signals to the hand muscles serving individual fingers (uni-digit muscles). The network was validated using three different training sets. Single digit muscles contributed from 25% to 50% of the total finger force. The master matrix and the enslaving matrix were computed; they characterize the ability of a given finger to enslave other fingers and its ability to be enslaved. Overall, the neural network modeling suggests that no direct correspondence exists between neural command to an individual finger and finger force. To produce a desired finger force, a command sent to an intended finger should be scaled in accordance with the commands sent to the other fingers.
Adult↗
Modulated cortical control of individual fingers in experienced musicians: an EEG study. Electroencephalographic study.
OBJECTIVE: The present research was designed to address the nature of interdependency between fingers during force production tasks in subjects with varying experience in performing independent finger manipulation. Specifically, behavioral and electroencephalographic (EEG) measures associated with controllability of the most enslaved (ring) and the least enslaved (index) fingers was examined in musicians and non-musicians. METHODS: Six piano players and 6 age-matched control subjects performed a series of isometric force production tasks with the index and ring fingers. Subjects produced 3 different force levels with either their index or ring fingers. We measured the isometric force output produced by all 4 fingers (index, ring, middle and little), including both ramp and static phases of force production. We applied time-domain averaging of EEG single trials in order to extract 4 components of the movement-related cortical potentials (MRCP) preceding and accompanying force responses. RESULTS: Three behavioral findings were observed. First, musicians were more accurate than non-musicians at reaching the desired force level. Second, musicians showed less enslaving as compared to non-musicians. And third, the amount of enslaving increased with the increment of nominal force levels regardless of whether the index or ring finger was used as the master finger. In terms of EEG measures, we found differences between tasks performed with the index and ring fingers in non-musicians. For musicians, we found larger MRCP amplitudes at most electrode sites for the ring finger. CONCLUSIONS: Our data extends previous enslaving research and suggest an important role for previous experience in terms of the independent use of the fingers. Given that a variety of previous work has shown finger independence to be reflected in cortical representation in the brain and our findings of MRCP amplitude associated with greater independence of fingers in musicians, this suggests that what has been considered to be stable constraints in terms of finger movements can be modulated by experience. SIGNIFICANCE: This work supports the idea that experience is associated with changes in behavioral and EEG correlates of task performance and may have clinical implications in disorders such as stroke or focal hand dystonia. Practice-related procedures offer useful approaches to rehabilitation strategies.
Adult↗
Genomic reduction and evolution of novel genetic membranes and protein-targeting machinery in eukaryote-eukaryote chimaeras (meta-algae).
Chloroplasts originated just once, from cyanobacteria enslaved by a biciliate protozoan to form the plant kingdom (green plants, red and glaucophyte algae), but subsequently, were laterally transferred to other lineages to form eukaryote-eukaryote chimaeras or meta-algae. This process of secondary symbiogenesis (permanent merger of two phylogenetically distinct eukaryote cells) has left remarkable traces of its evolutionary role in the more complex topology of the membranes surrounding all non-plant (meta-algal) chloroplasts. It took place twice, soon after green and red algae diverged over 550 Myr ago to form two independent major branches of the eukaryotic tree (chromalveolates and cabozoa), comprising both meta-algae and numerous secondarily non-photosynthetic lineages. In both cases, enslavement probably began by evolving a novel targeting of endomembrane vesicles to the perialgal vacuole to implant host porter proteins for extracting photosynthate. Chromalveolates arose by such enslavement of a unicellular red alga and evolution of chlorophyll c to form the kingdom Chromista and protozoan infrakingdom Alveolata, which diverged from the ancestral chromalveolate chimaera. Cabozoa arose when the common ancestor of euglenoids and cercozoan chlorarachnean algae enslaved a tetraphyte green alga with chlorophyll a and b. I suggest that in cabozoa the endomembrane vesicles originally budded from the Golgi, whereas in chromalveolates they budded from the endoplasmic reticulum (ER) independently of Golgi-targeted vesicles, presenting a potentially novel target for drugs against alveolate Sporozoa such as malaria parasites and Toxoplasma. These hypothetical ER-derived vesicles mediated fusion of the perialgal vacuole and rough ER (RER) in the ancestral chromist, placing the former red alga within the RER lumen. Subsequently, this chimaera diverged to form cryptomonads, which retained the red algal nucleus as a nucleomorph (NM) with approximately 464 protein-coding genes (30 encoding plastid proteins) and a red or blue phycobiliprotein antenna pigment, and the chromobiotes (heterokonts and haptophytes), which lost phycobilins and evolved the brown carotenoid fucoxanthin that colours brown seaweeds, diatoms and haptophytes. Chromobiotes transferred the 30 genes to the nucleus and lost the NM genome and nuclear-pore complexes, but retained its membrane as the periplastid reticulum (PPR), putatively the phospholipid factory of the periplastid space (former algal cytoplasm), as did the ancestral alveolate independently. The chlorarachnean NM has three minute chromosomes bearing approximately 300 genes riddled with pygmy introns. I propose that the periplastid membrane (PPM, the former algal plasma membrane) of chromalveolates, and possibly chlorarachneans, grows by fusion of vesicles emanating from the NM envelope or PPR. Dinoflagellates and euglenoids independently lost the PPM and PPR (after diverging from Sporozoa and chlorarachneans, respectively) and evolved triple chloroplast envelopes comprising the original plant double envelope and an extra outermost membrane, the EM, derived from the perialgal vacuole. In all metaalgae most chloroplast proteins are coded by nuclear genes and enter the chloroplast by using bipartite targeting sequences--an upstream signal sequence for entering the ER and a downstream chloroplast transit sequence. I present a new theory for the four-fold diversification of the chloroplast OM protein translocon following its insertion into the PPM to facilitate protein translocation across it (of both periplastid and plastid proteins). I discuss evidence from genome sequencing and other sources on the contrasting modes of protein targeting, cellular integration, and evolution of these two major lineages of eukaryote "cells within cells". They also provide powerful evidence for natural selection's effectiveness in eliminating most functionless DNA and therefore of a universally useful non-genic function for nuclear non-coding DNA, i.e. most DNA in the biosphere, and dramatic examples of genomic reduction. I briefly argue that chloroplast replacement in dinoflagellates, which happened at least twice, may have been evolutionarily easier than secondary symbiogenesis because parts of the chromalveolate protein-targeting machinery could have helped enslave the foreign plastids.
Chimera↗
Finger interaction during multi-finger tasks involving finger addition and removal.
The purpose of this study was to investigate changes in finger forces and their interactions in one-hand multi-finger force production tasks involving finger addition and removal. Eight male subjects were instructed to produce maximal (MVC) forces with explicitly instructed ('master') fingers. After reaching maximal force with a set of master fingers, the subjects added/removed one master finger while continuing to produce the MVC with the new group of master fingers. The non-instructed ('slave') fingers also produced forces (enslaving). Finger addition/removal led to changes in the forces of individual master fingers expected from earlier studies of maximal force production by different finger groups acting synchronously. A significant increase in the forces of remaining master fingers was observed after finger removal and a close-to-significant drop in the forces of previously recruited master fingers was observed after finger addition. These effects were larger when subjects started the task with a smaller number of explicitly involved fingers. The enslaving effects increased after finger addition while they did not change after finger removal. Forces produced by the same group of master fingers acting in different tasks showed no history-dependent effects. However, significant effects of history were seen in enslaving. These observations speak against hypotheses of more independent behaviour of effectors during their asynchronous involvement. They show that finger interaction may show effects of the history of finger involvement in a task.
Adult↗
Central mechanisms of finger interaction during one- and two-hand force production at distal and proximal phalanges.
In this study we used changes in the relative involvement of different muscle groups during force production at the distal (DT) and proximal (PR) phalanges to test and modify a hypothesis on the central organization of multi-finger control for tasks involving non-homologous elements in the two hands. Ten subjects produced maximal force with different finger combinations. Two symmetrical (PR/PR and DT/DT) and two asymmetrical (PR/DT and DT/PR) combinations of force application sites in the two hands were used. During one-hand tasks, higher forces were produced at the PR site. In multi-finger tasks, total peak force was smaller than the sum of peak forces in single-finger tasks by the involved fingers (force deficit). Force production by some fingers of a hand was accompanied by involuntary force production by other fingers (enslaving). Force deficit and enslaving were both higher at the PR site. Two-hand tasks were accompanied by an additional drop in the force of individual fingers, i.e., bilateral deficit (BD). When symmetrical sites of force production were used in the two hands, BD was lower for symmetrical finger groups than for asymmetrical groups. During tests at asymmetrical sites, BD was higher and did not depend on symmetry of involved finger groups. We conclude that within-a-hand force deficit and enslaving are likely to be of a central, neural origin. An earlier introduced hypothesis has been expanded assuming that excitatory projections to contralateral finger representations exist only for homologous elements (sub-synergies) of a multi-finger force production synergy, while only inhibitory projections connect non-homologous elements.
Adult↗
Age effects on force produced by intrinsic and extrinsic hand muscles and finger interaction during MVC tasks.
Finger-pressing forces are produced by activation of the intrinsic hand muscles, which are finger specific, and the extrinsic muscles that connect to multiple fingers. We tested a hypothesis of greater weakening of intrinsic hand muscles with age and quantified associated indexes of finger interaction such as enslaving (force production by unintended fingers) and force deficit (loss of finger force in multifinger tasks compared with single-finger tasks). Twelve young (23-35 yr old) and 12 elderly (70-95 yr old) men and women performed single-finger and four-finger maximal pressing tasks, in which force was applied at the proximal phalanges (PP, the intrinsic muscles are major focal force generators) and at the distal phalanges (DP, the extrinsic muscles are focal force generators). The decline in the peak force with age was greater at PP (30%) than at DP (19%). Larger indexes of finger interaction were observed at PP (enslaving = 17.2 +/- 9.4%, force deficit = 36.1 +/- 11.1%) than at DP (enslaving = 14.9 +/- 8.8%, force deficit = 27.7 +/- 10.8%) across ages and genders. We conclude that intrinsic hand muscles show disproportionate weakening with age. The greater indexes of finger interaction in PP tests with greater involvement of intrinsic hand muscles suggest that the finger interactions are predominantly of a central origin across ages and genders.
Adult↗
Finger coordination in persons with Down syndrome: atypical patterns of coordination and the effects of practice.
The study addresses an issue of possible relations between the apparent "clumsiness" of persons with Down syndrome (DS) and changes in indices of finger coordination. We hypothesized that persons with DS would prefer less challenging, safer motor strategies reflected in finger coordination patterns. Maximal single- and multi-finger force production (MVC) tasks and multi-finger tasks that required the production of a controlled time pattern (ramp) of total force were studied. As compared to typical persons, persons with DS showed lower peak forces, lower force deficit (loss of finger force in multi-finger tasks as compared to single-finger tasks), and higher enslaving (involuntary force production by fingers that are not required to produce force). They showed higher variance of total force computed across several trials for ramp tasks. Their total force variance was higher than the sum of the variances of individual finger forces over the ramp duration, while in control participants the relation was opposite during the middle and late thirds of the ramp. Persons with DS practiced force production tasks over 3 days, one group practicing only one of the tasks (the ramp task with all four fingers acting together) while the other group practiced MVC and ramp tasks (variable practice). Practice led to an increase in MVC, force deficit, and enslaving. The relation between the total force variance and the sum of the variances of individual finger forces became closer to the one observed in typical persons. The effects of practice were more pronounced in the variable practice group. We conclude that persons with DS have a deficit in control of both single fingers and multi-finger groups. They use a less challenging, suboptimal strategy of multi-finger coordination which does not take advantage of the possibility of error compensation among the fingers. Practice is an effective way of improving finger coordination in DS, particularly when using variable tasks.
Adolescent↗
Is the thumb a fifth finger? A study of digit interaction during force production tasks.
We studied indices of digit interaction in single- and multi-digit maximal voluntary contraction (MVC) tests when the thumb acted either in parallel or in opposition to the fingers. The peak force produced by the thumb was much higher when the thumb acted in opposition to the fingers and its share of the total force in the five-digit MVC test increased dramatically. The fingers showed relatively similar peak forces and unchanged sharing patterns in the four-finger MVC task when the thumb acted in parallel and in opposition to the fingers. Enslaving during one-digit tasks showed relatively mild differences between the two conditions, while the differences became large when enslaving was quantified for multi-digit tasks. Force deficit was pronounced when the thumb acted in parallel to the fingers; it showed a monotonic increase with the number of explicitly involved digits up to four digits and then a drop when all five digits were involved. Force deficit all but disappeared when the thumb acted in opposition to the fingers. However, for both thumb positions, indices of digit interaction were similar for groups of digits that did or did not include the thumb. These results suggest that, given a certain hand configuration, the central nervous system treats the thumb as a fifth finger. They provide strong support for the hypothesis that indices of digit interaction reflect neural factors, not the peripheral design of the hand. An earlier formal model was able to account for the data when the thumb acted in parallel to the fingers. However, it failed for the data with the thumb acting in opposition to the fingers.
Adult↗
Force and torque production in static multifinger prehension: biomechanics and control. II. Control.
The coordination of digits during combined force/torque production tasks was further studied using the data presented in the companion paper [Zatsiorsky et al. Biol Cybern this issue, Part I]. Optimization was performed using as criteria the cubic norms of (a) finger forces, (b) finger forces normalized with respect to the maximal forces measured in single-finger tasks, (c) finger forces normalized with respect to the maximal forces measured in a four-finger task, and (d) finger forces normalized with respect to the maximal moments that can be generated by the fingers. All four criteria failed to predict antagonist finger moments when these moments were not imposed by the task mechanics. Reconstruction of neural commands: The vector of neural commands c was reconstructed from the equation c=W(-1)F, where W is the finger interconnection weight matrix and F is the vector of finger forces. The neural commands ranged from zero (no voluntary force production) to one (maximal voluntary contraction). For fingers producing moments counteracting the external torque ('agonist' fingers), the intensity of the neural commands was well correlated with the relative finger forces normalized to the maximal forces in a four-finger task. When fingers produced moments in the direction of the external torque ('antagonist' fingers), the relative finger forces were always larger than those expected from the intensity of the corresponding neural commands. The individual finger forces were decomposed into forces due to 'direct' commands and forces induced by enslaving effects. Optimization of the neural commands resulted in the best correspondence between actual and predicted finger forces. The antagonist moments are, at least in part, due to enslaving effects: strong commands to agonist fingers also activated antagonist fingers.
Biomechanical Phenomena↗
Matrix analyses of interaction among fingers in static force production tasks.
The fingers on a hand show interactions in force production tasks. The interfinger connection matrices (IFMs) quantify these interactions (Li et al. 2002; Zatsiorsky et al. 2002b; Danion et al. 2003). The goal of the present study was to explore the differences in the IFMs of individual subjects and, in particular, to establish a procedure that may be used in the future for diagnostic purposes. Subjects ( n=20) pressed downward maximally with ten different combinations of the four fingers, index (I), middle (M), ring (R), and little (L): I, M, R, L, IM, MR, RL, IMR, MRL, and IMRL. Voluntary activation of a subset of the four fingers was accompanied by an involuntary force production by fingers that were not intentionally activated (enslaving). Interfinger connection matrices were computed for each subject by the artificial neural network. The similarities/dissimilarities (proximities) between the individual matrices were determined. This procedure was performed twice: (a) for nonnormalized IFMs whose elements represented the amount of force (in newtons) exerted by a finger i in response to a unit command to a finger j; and (b) for normalized IFMs, after dividing the elements of each IFM by the total force produced by the four fingers acting together (the elements of the matrix are in percents). The 20x20 matrix of the proximities was subjected to multidimensional scaling (MDS) to reduce the number of dimensions and identify the major ones. To interpret the meaning of the computed dimensions, they were regressed on a set of finger force parameters described in the text. For the nonnormalized IFMs an interpretable dimension was the strength of the subjects. For the normalized IFMs two dimensions were interpreted: (a) the location of the point of resultant force application along the mediolateral axis that is defined by the pattern of force sharing among the fingers and (b) the total contribution of the enslaved forces into the total finger force. We speculate that the similarity of typical everyday tasks across the population promotes the similarity of the IMFs reflecting optimal hand functioning over these tasks.
Adult↗
Indices of nonlinearity in finger force interaction.
Experiments with force production by subsets of fingers within the human hand have shown that finger interaction may be significantly nonlinear. In particular, this nonlinearity is reflected in the phenomenon of force deficit, a drop of the peak force of a finger when several fingers act simultaneously. We describe nonlinear effects in force relations within finger pairs, triplets, etc. Finger forces are represented as the sums of components resulting from force interactions within all subsets of the explicitly involved (master) fingers. The values of these components computed at extreme values of control signals, zero and unity, are taken as indices of such "elementary" force interactions. Indices of the first order reflect purposeful force production by a single master finger and its effects on forces produced by other fingers (enslaving). Indices of the second order reflect additional influences from pairs of simultaneously recruited master fingers, etc. Force interaction indices were computed based on finger forces measured in earlier experiments. Signs of indices alternated with their order, being positive for the indices of the first order (enslaving), negative for the indices of the second order (force deficit), positive for the indices of the third order, and mostly negative for the indices of the fourth order. Indices of the third and fourth orders reflect phenomena of force interaction not reported earlier. The study emphasizes the importance of nonlinear interactions among finger forces and introduces a set of independent indices that can be used to quantify such interactions in different subpopulations and their possible changes with practice and/or rehabilitation of the hand function.
Fingers↗
Catching and matching bars with different orientations.
The hypothesis that perception enslaves action is examined by assessing whether systematic distortions in perceptual judgments are reflected by inaccuracies in catching. In the first experiment, participants had to align manually the orientation of a reference bar placed at different distances in the frontoparallel plane. In the second experiment participants had to catch differently orientated moving bars, which became invisible at different distances from the interception point. In the matching experiment, systematic errors in the alignment of orientation were found in particular for oblique orientations, the magnitude of which increased with increasing distance of the reference bar. The inaccuracies in the final hand orientation during the catching task, however, did not mirror this pattern of deviations. The findings are interpreted to be more consistent with recent views that vision for perception (i.e., matching) and vision for action (i.e., catching) are dissociated than with the view that perception enslaves action.
Adult↗