Growth and gravitropism have been studied in three mutant strains of Arabidopsis thaliana L, that are resistant to auxin‐herbicide. Two of the mutations are allelic and recessive (aux‐1 and aux‐2) and are unlinked to a dominant mutation, Dwf, which confers a very high level of auxin‐resistance and is apparently lethal when homozygous. The aux‐1 and Dwf strains have altered response to gravity whereas aux‐2 appears to be gravitropically normal. After 96 h in the normal, vertical position only minor differences in elongation were observed between roots of wild‐type, aux‐1 and aux‐2, but the hypocotyls of aux‐1 were significantly retarded compared with the gravitropically normal aux‐2 and wild‐type. In the progeny of selfed Dwf plants, where both normal (dwf) and agravitropic (Dwf) seedlings are present, the Dwf seedlings had much longer roots and shorter hypocotyls than dwf+. During 22 h of continuous stimulation the optimum angle for gravitropism in wild‐type roots and hypocotyls was 135° (i.e. the organ points obliquely upwards), with decreasing responses in the order 90° and 45°. The agravitropic nature of the roots of aux‐1 was confirmed as no significant response was obtained at any of the stimulation angles. In marked contrast, the negative gravitropic response of aux‐1 hypocotyls was greater than the wild‐type response in terms of the final angle attained at 22 h, but between 6 and 22 h the elongation rate was lower in aux‐1. After varying stimulation periods in the horizontal position, the curvature which had developed, decreased rapidly and almost disappeared during ensuing rotation on clinostats (2 and 4 rpm). Rotation on the clinostats had no effect on the agravitropic behaviour of aux‐1.
The root anatomy and ultrastructure of the agravitropic Arobidopsis thaliana L. mutants Dwf and aux-1 were compared with the gravitropic mutant aux-2 and the wild type (WT) in an attempt to find an explanation for the lack of response to gravity. No differences were found in the organization of the root cap. The central part of the cap (columella) contains 5 storeys of developing, functioning and degenerating statocytes. Their ultrastructure is very similar in all four types of plant. Particular attention was paid to the distribution of rough endoplasmic reticulum (ER). Both in the WT and the mutants the ER is concentrated in the distal part at the "floor" of the cell. Light micrographs were used to compare the sedimentation rates of movable cell structures in normal and agravitropic root statocytes. A longitudinal movement of amyloplasts and nuclei was observed when the roots were inverted. In WT and aux-2 the rates were on average 6.3 micrometers h-1 (amyloplasts) and 2.1 micrometers h-1 (nucleus). In aux-1 the sedimentation rates were significantly lower: 2.4 and 0.6 micrometers h-1, respectively. Based on magnified electron micrographs of normal and inverted statocytes a morphometrical analysis of the distribution and redistribution of amyloplasts, nuclei, mitochondria, vacuoles and ER was made. The only significant difference was found in the redistribution of amyloplasts between aux-1 and the gravitropical normal types.
The pea mutant (Pisum sativum ageotropum) and the normal pea (P. sativum cv. Sabel) were compared in order to see if there were any differences in root anatomy or submorphology which could explain the presumed ageotropic behaviour of the mutant. In both types the root cap consists of a central core (columella) distinct from the peripheral part. The core contains five to six rows of columella cells, each consisting of 10 to 16 storeys of statocytes. The ultrastructure of the columella cells in the two types is very similar; the main difference is confined to the distribution of rough endoplasmic reticulum (ER), which in the mutant statocytes is evenly distributed throughout the cell, while in the normal pea statocytes it is mainly concentrated in the distal part at the “floor” of the cell. Using light micrographs, the movement of amyloplasts and nuclei have been followed in detail during a 40 min inversion period. The pattern of movement of the amyloplasts is apparently identical in the two types and the distances moved during the inversion period are 39 μm and 44 μm in the normal and mutant statocytes, respectively. The nucleus has not been observed to move in normal pea; a slight rearrangement of the nucleus position can be observed during the period 30 to 40 min after the start of inversion of the mutant. Based on magnified electron micrographs of the statocytes a morphometrical analysis was made of five cell structures – amyloplasts, nuclei, mitochondria, vacuoles and ER – which appeared to be freely movable or redistributable under the influence of the gravitational force.
In an attempt to explain the influence of gravity on the behaviour of ageotropic plant organs, a pea mutant (Pisum sativum ageotropum) and normal pea (Pisum sativum cv. Sabel) were examined. The mutant has a significantly lower germination rate (large seeds: 25%, small seeds: 10%) than normal pea seeds (55%). Removal of testa increased germination dramatically, the values obtained were 63 and 89%, respectively. Immediately after imbibition the mutant from which the testa had been removed, developed more slowly than normal pea seeds; after 28 h the difference in elongation rate between the two types was reversed. When continuously stimulated geotropically in the horizontal position the elongation in the mutant is larger than in the normal pea roots kept in the same position. During a 24 h period starting 48 h after imbibition the mutant root elongated 45.0 mm while the value for the normal pea root was 11.5 mm. The course of the geotropic curvature in roots of the two types has been followed during a period of 24 h. Normal pea roots develop an asymmetry in the extreme root tip region after 30 min of horizontal stimulation. After prolonged stimulation (exceeding 2 h) the asymmetry has disappeared and the curvature distributed over the entire growth region. When roots of normal pea are stimulated continuously at various angles, the optimum angle of geotropic response is 90° with decreasing responses in the order 135° (i.e. the root tip is pointing obliquely upward) and 45°. The presumed ageotropic behaviour of the mutant has only to a certain extent been confirmed in the present study. When stimulated at 135° a slight positive curvature developed; stimulation at 90° and 45° gave a slight negative curvature.
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