Movement of Amyloplasts in the Statocytes of Geotropically Stimulated Roots. The Pre-Inversion Effect

Previously inverted Lepidium roots were placed in a horizontal position and the amyloplasts in the statocytes of the root cap allowed to fall through their entire range of movement across the cell. Under these conditions the amyloplasts first follow a mainly downward course for 6 to 8 min at a speed between 0.5 and 0.8 μm per min. For the next 10 min they move slightly more slowly in a direction away from the apical end of the cell, still sinking somewhat, but without reaching the plasmalemma along the lower wall. Previous experiments have shown that conditions assumed to allow the amyloplasts to slide parallel to the longitudinal cell walls are those that give rise to the largest geotropic curvatures. Such conditions are for instance (1) stimulation at 135° (root tips pointing obliquely upward) and (2) inversion of roots for 16 min followed by stimulation at 45°. Treatments assumed not to permit extensive sliding of the amyloplasts produce smaller geotropic curvatures, namely (3) stimulation at 45° without pre-inversion and (4) inversion followed by stimulation at 135°. The location of the amyloplasts after these four kinds of treatment has now been determined on photomicrographs and the assumptions concerning the paths and extent of sliding of the amyloplasts confirmed. Observations on electron micrographs showed that under all conditions the amyloplasts are separated from the plasmalemma by other organelles, such as ER, nucleus or vacuoles. In roots rotated for 15 min parallel to the horizontal axis of the klinostat at 2 rpm, the amyloplasts are not clumped together as densely as in normal, inverted or stimulated roots, but they are not scattered over the entire cell volume. The statolith function of the amyloplasts is discussed in view of these and other observations.     

GEOTROPISM OF AGRIOLIMAX

On an inclined glass plate the slug Agriolimax orients and creeps upward or downward. The angle of orientation on the plane (θ) is proportional to the logarithm of the component of gravity in the creeping plane. The coefficient of variability of the measured values of (θ) decreases linearly as the logarithm at the gravity component in the creeping plane increases. The cosine of the angle of orientation decreases almost directly in proportion to the sine of the angle of inclination of the creeping plane to the horizontal, as previously found for young rats (Crozier and Pincus). But a more satisfactory formulation for the present case shows that the sine of the angle of orientation (θ) decreases in direct proportion to the increase of the reciprocal of the sine of the angle of inclination of the creeping plane. This formulation is derived from the theory that the geotropic orientation is limited by the threshold difference between the pull of the body mass on the mutually inclined longitudinal muscles at the anterior end of the slug.     

水稻无侧根突变体的根向重力性异常

用化学诱变剂 (NaN3 )处理粳稻品种大力 (O ryzasativaL .cv .Oochikara) ,得到具有 2 ,4 D抗性、无侧根和根向重力性异常的突变体RM 10 9。对原品种为父本和突变体为母本的杂交后代F1、F2 根向重力性的遗传分离进行了研究。结果表明 :突变体的根向重力性异常 ,其性状是单显性基因控制且不受光照和黑暗培养的影响。通过对根冠组织切片观察发现 :突变体根冠中含淀粉体的细胞数量比大力少 ,根冠细胞中淀粉体的直径为原品种的 5 0 %且集中排列于细胞内的一角 ,其排列沉积方向与重力方向相同。推测 :突变体的根向重力性异常与淀粉体直径变小有关     

ON THE EQUILIBRATION OF GEOTROPIC AND PHOTOTROPIC EXCITATIONS IN THE RAT

The intensity of light required to just counterbalance geotropic orientation of young rats, with eyelids unopened, is so related to the angle of inclination (α) of the creeping plane that the ratio log I/log sin α is constant. This relationship, and the statistical variability of I as measured at each value of α, may be deduced from the known phototropic and the geotropic conduct as studied separately, and affords proof that in the compounding of the two kinds of excitation the rat is behaving as a machine.     

GEOTROPIC EXCITATION IN HELIX

Rotation of an inclined surface on which Helix is creeping straight upward, such that the axis of the animal is turned at a right angle to its previous position, but in the same plane, leads to negatively geotropic orientation after a measurable latent period or reaction time. The duration of the latent period is a function of the slope of the surface. The magnitude of the standard deviation of the mean latent period is directly proportional to the mean latent period itself, so that the relative variability of response is constant. The dependence of reaction time upon extent of displacement from symmetrical orientation in the gravitational field is found also by tilting the supporting surface, without rotation in the animal''s own plane. On slopes up to 55°, the relation between latent period and the sine of the slope is hyperbolic; above this inclination, the latent period sharply declines. This change in the curve is not affected by the attachment of moderate loads to the snail''s shell (up to 1/3 of its own mass), and is probably a consequence of loss of passive stable equilibrium when rotated. When added loads do not too greatly extend the snail''s anterior musculature, the latent period for the geotropic reaction is decreased, and, proportionately, its σ. These facts are discussed from the standpoint that geotropic excitation in these gasteropods is due to impressed muscle-tensions.     

Recovery of Geotropism after Removal of the Root Cap

Removal of the cap from the primary roots of Zea mays and Triticumaestivum renders the roots unresponsive to gravity. In bothspecies a geotropic response is recovered before a new cap hasstarted to regenerate. Immediately after decapping amyloplastsstart to develop in cells of the root apex and it is proposedthat as the development of amyloplasts continues so they becomefunctional as gravity sensors. It is also suggested that theamyloplasts may be the source of an inhibitor that has beenpostulated to be the intermediary between the perception ofgravity and the geotropic response.     

Gravity perception in decapped roots of Zea mays

Time-lapse photography and light microscopy were used to determine whether or not sedimentation of the newly developed amyloplasts in the apex of Zea mays L. roots occurred at the time when geotropic responsiveness reappears following removal of the cap. All decapped roots exhibiting a geotropic response had some amyloplast sedimentation in the apical cortical cells. Exposing decapped roots to a centrifugal acceleration of 25 g for 4 h showed that amyloplasts of a similar size and development were not displaced within the cytoplasm when this treatment began 12 h after decapping, whereas displacement did occur when the treatment began 24 h after decapping. This finding indicates the occurrence of a change in the physical characteristics of the cytoplasm between 12 h and 24 h after removing of the cap, which allows amyloplast movement and thus restores gravity perception.     

Geoperception in primary and lateral roots of Phaseolus vulgaris (Fabaceae). I. Structure of columella cells

Primary roots of Phaseolus vulgaris (Fabaceae) are positively geotropic, while lateral roots are not responsive to gravity In order to elucidate the structural basis for this differential georesponse, we have performed a qualitative and quantitative analysis of the ultrastructure of columella cells of primary and lateral roots of P. vulgaris. Root systems were fixed in situ so as not to disturb the ultrastructure of the columella cells. The columellas of primary roots are more extensive than those of lateral roots. The volumes of columella cells of primary roots are approximately twice those of columella cells of lateral roots. However, columella cells of primary roots contain greater absolute volumes and numbers of all cellular components examined than do columella cells of lateral roots. Also, the relative volumes of cellular components in columella cells of primary and lateral roots are statistically indistinguishable. The endoplasmic reticulum is sparse and distributed randomly in both types of columella cells. Both types of columella cells contain numerous sedimented amyloplasts, none of which contact the cell wall or form complexes with other cellular organelles. Therefore, positive geotropism by roots must be due to a factor(s) other than the presence of sedimented amyloplasts alone. Furthermore, it is unlikely that amyloplasts and plasmodesmata form a multi-valve system that controls the movement of growth regulating substances through the root cap.     

GEOTROPISM AND MUSCLE TENSION IN HELIX

1. The snail Helix aspersa Müller, is negatively geotropic during the daytime, but positive or indifferent at night. 2. The precision of geotropic orientation is a function of the gravity component acting on the body. 3. The rate of geotropic locomotion is also determined by the gravity component (sine of the angle of inclination). 4. The rate of upward movement is increased 1.51 times at 45° inclination by loading the snail with one-half its weight. No such increase is seen in loaded snails creeping on a horizontal surface. 5. Moderate centrifugation results in orientation and locomotion towards the center of rotation. 6. A response analogous to the homostrophic reflex occurs when a backward pull to right or to left is exerted on the shell. Bilaterally equal tension applied to the shell causes locomotion along a path parallel and opposite to the direction of the pull. 7. All the observations go to show that the stimulus for geotropic orientation and locomotion is tension of the body muscles produced by the downward pull of gravity, and that the stimulus is received by the proprioreceptors of these muscles. Otolith apparatus and analogous organs, when present, may assist in the response, but they do not seem to be requisite in all cases. Since the precision of orientation and the rate of locomotion are functions of the gravity component acting on the body, the muscle tension theory of the geotropic reactions accords fully with Loeb''s tropism doctrine for animals.     

Geotropism and the lateral transport of auxin in the corn mutant amylomaize

Summary In coleoptiles of the amylomaize corn mutant (AM), the amyloplasts are much reduced in size in comparison with the wild type corn (WT), permitting a comparison of geotropic responsiveness as related to lateral displacement of amyloplasts and lateral transport of auxin. The amyloplasts of AM showed 30–40% lesser lateral redistribution in response to horizontal exposure in comparison with WT. With geotropic stimulation, the lateral transport of auxin in the direction of growth was 40–80% less, and the geotropic curvature by the coleoptiles was also significantly less in the mutant as compared with WT. These correlations support the hypothesis that the starch plastids serve as gravity sensors in the geotropic responses of coleoptiles.     

Relation entre réaction géotropique et évolution du statenchyme dans la racine d'Asperge

Relationship between the Geotropic Response and the Evolution of the Statenchyma in Roots of Asparagus officinalis. The evolution of the statenchyma in roots of Asparagus of ficinalis seedlings, grown in obscurity, was followed during the first 17 days. After 7 days of etiolation, a decrease of both the average diameter of the amyloplasts and the average number of these organelles was observed in the central root cap cells. If the seedlings were illuminated (with white light) from the 7th day, the average number of statoliths increased rapidly in the statocytes. The volume of these organelles undergoes the same variation in etiolated and in illuminated plants. The initial rate of curvature (V_i) of the roots (stimulated in a horizontal position) and the volume of amyloplasts (V_ac) in their caps were analysed as a function of the time of germination in obscurity (from the 8th to the 17th day). It was found that V_i increased as a linear function of the logarithm of V_ac, which confirms that the weight of the amyloplasts of the statocytes may play a role in the geotropic stimulation of the roots.     

The Mechanism of Geoperception in Lentil Roots

The sediment of the amyloplasts in the statocytes of lentilroots was analysed in two different samples of seedlings. Theroots of the first sample (HP) were grown in a horizatal position,whereas thoes of the second sample (VP+20) were grown in a horizontalposition and then exposure to give a response). It is demonstratedthat the statolish can almost enter into contact with the plasmamembrane lining the longitudinal wall of the statocytes of theHP sample. However, these organelles in the VP+20 sample areable to provoke a geotropic stimulation though they are sedimentedat a distance of 0.1350–0.2600 µm from the plasmalemma.From these data it is concluded that the cytoplasmic structureswhich may play a role in the geotropic stimulation are: (1)the endoplasmic reticulum located along the longitudianl wallof the statocytes and (2) the microtubules or the plasmalemmaif the action of the statoliths is transmitted by the parietalcytoplasm. The large aggregations of endoplasmic reticulum whichare situated in the distal part of the central root cap cellswould not have any role in the perception of gravity by roots.These results are discussed in the light of recent work showingthe action of a growth inibitor in the geotropic reaction ofroots.     

THE GEOTROPIC RESPONSE IN ASTERINA

Upon a surface inclined at angle α Asterina gibbosa orients upward during negatively geotropic creeping until the average angle (θ) of the path is such that Δ sin θ/Δ sin α = const. This is true also in positively geotropic movement. The direction of orientation may be temporarily reversed by mechanical disturbance. The variation of θ is greater at low slopes. Tests with directed impressed pulls, due to an attached cork float, show that the pull upon the tube feet is of primary consequence for the determination of θ. When the component of gravitational pull in the direction of movement reaches a fraction of the total pull which is proportional to the gravitational vector parallel to the surface, the laterally acting component is ineffective. On this basis, it follows that Δ sin θ/Δ sin α = const.     

GEOTROPIC ORIENTATION IN ARTHROPODS : I. MALACOSOMA LARV?.

The geotropic orientation of caterpillars of Malacosoma americana during progression upon a surface inclined at angle α to the horizontal is such that the path makes an average angle θ upward on the plane, of a magnitude proportional to log sin α. More precisely, the product (sin α) (sin θ) is constant. This is traced to the fluctuation of the pull of the head region upon the lateral musculature of the upper side during the side to side swinging implicated in progression.     

The Optimum Angle of Geotropic Stimulation and its Relation to the Starch Statolith Hypothesis

Roots which are turned from their normal direction to directions at various angles with the plumb line develop the largest geotropic curvatures during a subsequent klinostat rotation period when the stimulation angle is well above the horizontal. In experiments with roots of Lepidium sativum L., the optimum is located at 120 to 140° when the stimulation time is between 2 and 15 min. If this fact is to be explained by the movements of amyloplasts in the root cap cells, one would expect roots which bad been kept inverted before the stimulation (so that the moveable amyloplasts are accumulated in the opposite end of the cells) to show an optimum angle well below 90°. — Pre-inversion of the roots did suppress the curvatures produced by stimulation at angles larger than 90° when measured after 10 to 30 min of klinostat rotation. This suppression may be taken as a support for the starch statolith hypothesis, since the amyloplasts in pre-inverted roots placed at angles exceeding 90° have a restricted opportunity to slide along the cell walls compared to non-inverted roots placed at the same angles. In pre-inverted roots measured after a period of klinostat rotation, however, no optimum was found at angles below 90°. When the stimulation time was 3.75 min, the response curves were nearly symmetrical about 90°. Stimulation for 15 min, on the other hand, resulted in curvatures which were much larger (although suppressed in comparison with non-inverted roots) when the stimulation angle was 165° than when it was 15°. During the 15 min stimulation period itself, however, pre-inverted roots curved 0.3° when stimulated at 15, but only 3.4° at 165°. This small difference was very highly significant and is in agreement with the starch statolith hypothesis insofar as the amyloplasts in pre-inverted roots placed at 15° have the greatest opportunity to slide along the cell walls. The lack of further development (and the actual decrease) of their curvatures during the subsequent klinostat rotation must then be due to other, depressing, factors, summarily designated as tonic. At angles above 90°, the tonic factors are either absent or even enhancing. Tbe tonic effects cannot be explained by amyloplast movements.     

Development of the root crown in some conifers

Summary Large and small stumps of 3 spruce species, a pine, a fir and hemlock were pulled up from 15 year plantations. Roots originating from the root crown were counted (<0.5 cm) or, if larger, their diameters were measured, taper, eccentricity, compass bearing, inclination, position on the rootstock and basal age (by annual rings). The sequence of events leading to the developed skeletal root system was reconstructed. Hypotheses to account for the distinct populations of roots were examined. In all species, major roots originated between 2 and 7 yrs after planting and grew thicker and more eccentric at their bases than minor roots of the same age. Positively geotropic roots were distinct from plagiogeotropic roots.     

Reversible effects of green and orange-red radiation on plant cell elongation

The differential cell elongation of cress (Lepidium sativum) roots that results in geotropic bending can be decreased by green radiation with an action spectrum peaking at 550 nm. This decrease can be negated by prior or simultaneous irradiation by orange-red light with an action spectrum peaking at 620 nm. The green radiation appears to be effective during the cell elongation phase of geotropic response.     

The T cell receptor V alpha 11 gene family. Analysis of allelic sequence polymorphism and demonstration of J alpha region-dependent recognition by allele-specific antibodies.

Allelic polymorphism in TCR loci may play an important role in shaping the T cell repertoire and in disease susceptibility. We have used a combination of antibody and sequence analysis to investigate polymorphism in the murine V alpha 11 family. Two different antibodies have been analyzed that recognize particular V alpha 11 family members of the V alpha b and V alpha d haplotypes. One antibody shows J alpha dependency, suggesting a conformational element to the epitope. Investigation of the anti-V alpha 11 staining pattern on different mouse strains indicates that there is a marked influence of MHC haplotype on V alpha 11 selection and that V alpha 11 is preferentially expressed on CD4+ cells. Sequence analysis of V alpha 11 genes from the V alpha a, V alpha b, and V alpha d haplotypes shows two potential regions for the haplotype-specific epitopes. The relatedness of the different V alpha 11 family members from different haplotypes suggests that the V alpha 11.1/11.2 gene duplication is relatively recent, but that V alpha 11.3 separated much earlier. Differences between V alpha 11.3 and V alpha 11.1/11.2 are concentrated in the putative complementarity determining regions (CDR), whereas differences between alleles are not clearly clustered. However, the V alpha 11.1a and V alpha 11.1d alleles differ from V alpha 11.1b and V alpha 11.2b in CDR1. A V alpha 11.2-expressing anti-cytochrome c T cell has the same V-J junction as a V alpha 11.1-bearing cell with a similar fine specificity, indicating that V alpha 11.1b and V alpha 11.2b do not contribute different Ag specificities.     

An explanation of the inhibition of root growth caused by indole-3-acetic Acid

Low concentrations of indole-3-acetic acid inhibit the growth of pea root sections by inducing the formation of the growth regulator, ethylene gas. Ethylene is produced within 15 to 30 minutes after indole-3-acetic acid is applied and roots begin to swell immediately after they are exposed to the gas. Carbon dioxide competitively inhibits ethylene action in roots, impedes their geotropic response, and partially reinstates auxin inhibited growth. It is concluded that ethylene participates in the geotropic response of roots, but not that of stems.     

GEOTROPIC ORIENTATION IN ARTHROPODS : III. THE FIDDLER CRAB UCA.

On an inclined surface the fiddler crab Uca pugnax, during sidewise progression, orients upward through an angle θ on the surface. The extent of negatively geotropic orientation (θ) is a rectilinear function of sin α, where α is the inclination of the surface to the horizontal. This equation differs from that describing the geotropic orientation of various other animals. The difference is traced to the fact that from an initial position with the transverse axis of the body horizontal the crab is required to turn upward to an extent such that the vertical line from its center of gravity pierces the inclined surface within the base of support provided by the legs. This leads to the equation sec θ/tan α = const., which is obeyed within the limits of precision of the measurements. This type of control of geotropic orientation represents an extension of the "muscle tension theory," and is in no sense in conflict with this view. The assumptions underlying the analytical expression connecting θ and α are verified by the asymmetry in the orientation of male fiddlers, which is shown to be due to the presence of the enlarged chela and which disappears when the claws are removed.