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Keywords: Auxin; gravitropism; hydrotropism; phototropism; roots EN Auxin gravitropism hydrotropism phototropism roots 4892 4895 4 09/15/23 20230913 NES 230913 When exposed to one-sided blue light, the roots of plants such as I Arabidopsis thaliana i exhibit negative phototropism and curve away from the light. Transgenic I GNOMpro:GNOM-GFP i roots showed perfect phototropism, identical to wild-type roots, but transgenic I SMBpro:GNOM-GFP i and I SCRpro:GNOM-GFP i roots showed reduced phototropism. Overall, the study shows that MIZ1 acts in the cortex of the Arabidopsis root elongation zone not only in hydrotropism but also in phototropism (Fig. [12] found out that the same tissues in Arabidopsis roots involved in MIZ1- and GNOM/MIZ2-regulated hydrotropism are also required for phototropism. [Extracted from the article]

In response to unilateral blue light illumination, roots of some plant species such as Arabidopsis thaliana exhibit negative phototropism (bending away from light), which is important for light avoidance in nature. MIZU-KUSSEI1 (MIZ1) and GNOM/MIZ2 are essential for positive hydrotropism (i.e. in the presence of a moisture gradient, root bending towards greater water availability). Intriguingly, mutations in these genes also cause a substantial reduction in phototropism. Here, we examined whether the same tissue-specific sites of expression required for MIZ1- and GNOM/MIZ2-regulated hydrotropism in Arabidopsis roots are also required for phototropism. The attenuated phototropic response of miz1 roots was completely restored when a functional MIZ1–green fluorescent protein (GFP) fusion was expressed in the cortex of the root elongation zone but not in other tissues such as root cap, meristem, epidermis, or endodermis. The hydrotropic defect and reduced phototropism of miz2 roots were restored by GNOM/MIZ2 expression in either the epidermis, cortex, or stele, but not in the root cap or endodermis. Thus, the sites in root tissues that are involved in the regulation of MIZ1- and GNOM/MIZ2-dependent hydrotropism also regulate phototropism. These results suggest that MIZ1- and GNOM/MIZ2-mediated pathways are, at least in part, shared by hydrotropic and phototropic responses in Arabidopsis roots. [ABSTRACT FROM AUTHOR]

Hydrotropism is the directed growth of roots toward the water found in the soil. However, mechanisms governing interactions between hydrotropism and gravitropism remain largely unclear. In this study, we found that an air system and an agar–sorbitol system induced only oblique water-potential gradients; an agar–glycerol system induced only vertical water-potential gradients; and a sand system established both oblique and vertical water-potential gradients. We employed obliquely oriented and vertically oriented experimental systems to study hydrotropism in Arabidopsis and tomato plants. Comparative analyses using different hydrotropic systems showed that gravity hindered the ability of roots to search for obliquely oriented water, whilst facilitating roots' search for vertically oriented water. We found that the gravitropism-deficient mutant aux1 showed enhanced hydrotropism in the oblique orientation but impaired root elongation towards water in the vertical orientation. The miz1 mutant exhibited deficient hydrotropism in the oblique orientation but normal root elongation towards water in the vertical orientation. Importantly, in contrast to miz1 , the miz1 / aux1 double mutant exhibited hydrotropic bending in the oblique orientation and attenuated root elongation towards water in the vertical orientation. Our results suggest that gravitropism is required for MIZ1-regulated root hydrotropism in both the oblique orientation and the vertical orientation, providing further insight into the role of gravity in root hydrotropism. [ABSTRACT FROM AUTHOR]

Fresh water is an increasingly scarce resource for agriculture. Plant roots mediate water uptake from the soil and have developed a number of adaptive traits such as hydrotropism to aid water foraging. Hydrotropism modifies root growth to respond to a water potential gradient in soil and grow towards areas with a higher moisture content. Abscisic acid (ABA) and a small number of genes, including those encoding ABA signal transducers, MIZ2/GNOM, and the hydrotropism-specific MIZ1, are known to be necessary for the response in Arabidopsis thaliana, whereas the role of auxin in hydrotropism appears to vary depending on the plant species. This review will describe recent progress characterizing the hormonal regulation of hydrotropism. Recent advances in identifying the sites of hydrotropic perception and response, together with its interaction with gravitropism, will also be discussed. Finally, I will describe putative mechanisms for perception of the water potential gradient and a potential role for hydrotropism in acclimatizing plants to drought conditions. [ABSTRACT FROM AUTHOR]

The direction of auxin transport changes in gravistimulated roots, causing auxin accumulation in the lower side of horizontally reoriented roots. This study found that auxin was similarly involved in hydrotropism and gravitropism in rice and pea roots, but hydrotropism in Lotus japonicus roots was independent of both auxin transport and response. Application of either auxin transport inhibitors or an auxin response inhibitor decreased both hydrotropism and gravitropism in rice roots, and reduced hydrotropism in pea roots. However, Lotus roots treated with these inhibitors showed reduced gravitropism but an unaltered or an enhanced hydrotropic response. Inhibiting auxin biosynthesis substantially reduced both tropisms in rice and Lotus roots. Removing the final 0.2 mm (including the root cap) from the root tip inhibited gravitropism but not hydrotropism in rice seedling roots. These results suggested that modes of auxin involvement in hydrotropism differed between plant species. In rice roots, although auxin transport and responses were required for both gravitropism and hydrotropism, the root cap was involved in the auxin regulation of gravitropism but not hydrotropism. Hydrotropism in Lotus roots, however, may be regulated by a novel mechanism that is independent of both auxin transport and the TIR1/AFBs auxin response pathway. [ABSTRACT FROM AUTHOR]

Background and Aims Root hydrotropism is a response to water-potential gradients that makes roots bend towards areas of higher water potential. The gene MIZU-KUSSEI1 (MIZ1) that is essential for hydrotropism in Arabidopsis roots has previously been identified. However, the role of root hydrotropism in plant growth and survival under natural conditions has not yet been proven. This study assessed how hydrotropic response contributes to drought avoidance in nature. Methods An experimental system was established for the study of Arabidopsis hydrotropism in soil. Characteristics of hydrotropism were analysed by comparing the responses of the miz1 mutant, transgenic plants overexpressing MIZ1 (MIZ1OE) and wild-type plants. Key Results Wild-type plants developed root systems in regions with higher water potential, whereas the roots of miz1 mutant plants did not show a similar response. This pattern of root distribution induced by hydrotropism was more pronounced in MIZ1OE plants than in wild-type plants. In addition, shoot biomass and the number of plants that survived under drought conditions were much greater in MIZ1OE plants. Conclusions These results show that hydrotropism plays an important role in root system development in soil and contributes to drought avoidance, which results in a greater yield and plant survival under water-limited conditions. The results also show that MIZ1 overexpression can be used for improving plant productivity in arid areas. [ABSTRACT FROM PUBLISHER]

Abscisic acid (ABA) signaling plays a critical role in regulating root growth and root system architecture. ABA-mediated growth promotion and root tropic response under water stress are key responses for plant survival under limiting water conditions. In this work, we have explored the role of Arabidopsis (Arabidopsis thaliana) PYRABACTIN RESISTANCE1 (PYR1)/PYR1-LIKE (PYL)/REGULATORY COMPONENTS OF ABA RECEPTORS for root ABA signaling. As a result, we discovered that PYL8 plays a nonredundant role for the regulation of root ABA sensitivity. Unexpectedly, given the multigenic nature and partial functional redundancy observed in the PYR/PYL family, the single pyl8 mutant showed reduced sensitivity to ABA-mediated root growth inhibition. This effect was due to the lack of PYL8-mediated inhibition of several clade A phosphatases type 2C (PP2Cs), since PYL8 interacted in vivo with at least five PP2Cs, namely HYPERSENSITIVE TO ABA1 (HAB1), HAB2, ABA-INSENSITIVE1 (ABI1), ABI2, and PP2CA/ABA-HYPERSENSITIVE GERMINATION3 as revealed by tandem affinity purification and mass spectrometry proteomic approaches. We also discovered that PYR/PYL receptors and clade A PP2Cs are crucial for the hydrotropic response that takes place to guide root growth far from regions with low water potential. Thus, an ABA-hypersensitive pp2c quadruple mutant showed enhanced hydrotropism, whereas an ABA-insensitive sextuple pyr/pyl mutant showed reduced hydrotropic response, indicating that ABA-dependent inhibition of PP2Cs by PYR/PYLs is required for the proper perception of a moisture gradient. [ABSTRACT FROM AUTHOR]

Because of their sessile nature, plants evolved several mechanisms to tolerate or avoid conditions where water is scarce. The molecular mechanisms contributing to drought tolerance have been studied extensively, whereas the molecular mechanism underlying drought avoidance is less understood despite its importance. Several lines of evidence showed that the roots sense the moisture gradient and grow toward the wet area: so-called hydrotropism. We previously identified MIZU-KUSSEI (MIZ) 1 and MIZ2/GNOM as genes responsible for this process. To gain new insight into the molecular mechanism of root hydrotropism, we generated overexpressors of MIZ1 (MIZ1OEs) and analyzed their hydrotropic response. MIZ1OEs had a remarkable enhancement of root hydrotropism. Furthermore, a greater number of MIZ1OE root cells remained viable under hydrostimulated conditions than those of the wild type, which might contribute to retaining root growth under hydrostimulated conditions. Although overexpression of MIZ1 also caused a slight decrease in the root gravitropic response, it was not attributable to the enhanced hydrotropic response. In addition, miz2 mutation or the auxin response inhibitor nullified the enhanced hydrotropic response in MIZ1OEs. Furthermore, the expression of MIZ1 did not alter the expression of typical genes involved in drought tolerance. These results suggest that MIZ1 positively regulates hydrotropism at an early stage and its overexpression results in an enhancement of signal transduction unique to root hydrotropism to increase the degree of hydrotropic root bending. [ABSTRACT FROM AUTHOR]

Roots are highly plastic and can acclimate to heterogeneous and stressful conditions. However, there is little knowledge of the effect of moisture gradients on the mechanisms controlling root growth orientation and branching, and how this mechanism may help plants to avoid drought responses. The aim of this study was to isolate mutants of Arabidopsis thaliana with altered hydrotropic responses. Here, altered hydrotropic response 1 (ahr1), a semi-dominant allele segregating as a single gene mutation, was characterized. ahr1 directed the growth of its primary root towards the source of higher water availability and developed an extensive root system over time. This phenotype was intensified in the presence of abscisic acid and was not observed if ahr1 seedlings were grown in a water stress medium without a water potential gradient. In normal growth conditions, primary root growth and root branching of ahr1 were indistinguishable from those of the wild type (wt). The altered hydrotropic growth of ahr1 roots was confirmed when the water-rich source was placed at an angle of 45° from the gravity vector. In this system, roots of ahr1 seedlings grew downward and did not display hydrotropism; however, in the presence of cytokinins, they exhibited hydrotropism like those of the wt, indicating that cytokinins play a critical role in root hydrotropism. The ahr1 mutant represents a valuable genetic resource for the study of the effects of cytokinins in the differential growth of hydrotropism and control of lateral root formation during the hydrotropic response. [ABSTRACT FROM PUBLISHER]

Plant organ development is important for adaptation to a changing environment. Genetic and physiological studies have revealed that plant hormones play key roles in lateral root formation. In this study, we show that MIZU-KUSSEI1 (MIZ1), which was identified originally as a regulator of hydrotropism, functions as a novel regulator of hormonally mediated lateral root development. Overexpression of MIZ1 (MIZ1OE) in roots resulted in a reduced number of lateral roots being formed; however, this defect could be recovered with the application of auxin. Indole-3-acetic acid quantification analyses showed that free indole-3-acetic acid levels decreased in MIZ1OE roots, which indicates that alteration of auxin level is critical for the inhibition of lateral root formation in MIZ1OE plants. In addition, MIZ1 negatively regulates cytokinin sensitivity on r development. Application of cytokinin strongly induced the localization of MIZl-green fluorescent protein to lateral root primordia, which suggests that the inhibition of lateral root development by MIZ1 occurs downstream of cytokinin signaling. Surprisingly, miz2, a weak allele of gnom, suppressed developmental defects in MIZ1OE plants. Taken together, these re suggest that MIZ1 plays a role in lateral root development by maintaining auxin levels and that its function requires GNOM activity. These data provide a molecular framework for auxin-dependent organ development in Arabidopsis (Arabidopsis thaliana). [ABSTRACT FROM AUTHOR]

Plants are sessile in nature, and need to detect and respond to many environmental cues in order to regulate their growth and orientation. Indeed, plants sense numerous environmental cues and respond via appropriate tropisms, and it is widely accepted that auxin plays an important role in these responses. Recent analyses using Arabidopsis have emphasized the importance of polar auxin transport and differential auxin responses to gravitropism. Even so, the involvement of auxin in hydrotropism remains unclear. To clarify whether or not auxin is involved in the hydrotropic response, Arabidopsis seedlings were treated with inhibitors of auxin influx (3-chloro-4-hydroxyphenylacetic acid), efflux (1-naphthylphthalemic acid and 2,3,5-triiodobenzoic acid), and response (p-chlorophenoxyisobutylacetic acid), and their effects were examined on both hydrotropic and gravitropic responses. In agreement with previous reports, gravitropism was inhibited by all the chemicals tested. By contrast, only an inhibitor of the auxin response (p-chlorophenoxyisobutylacetic acid) reduced hydrotropism, whereas inhibitors for influx or efflux of auxin had no effect. These results suggest that auxin response, apart from its polar transport, plays a definite role in hydrotropic response, and will evoke a new concept for the auxin-mediated regulation of tropisms. [ABSTRACT FROM PUBLISHER]

Presents a historical perspective analysis of osmogenetics. Systematic and definitive quantitative analyses of water absorption and movement within plants; Principles of chemical and water potential of osmosis; Emergence of the fundamental physical-chemical principles of plant water movement.

In the present study, root hydrotropism in an agravitropic mutant of Pisum sativum L. grown in vermiculite with a steep water potential gradient was examined. When wet and dry vermiculite were placed side by side, water diffused from the wet (-0·04 MPa) to the dry (-1·2 MPa) and a steep water potential gradient became apparent in the dry vermiculite close to the boundary between the two. The extent and location of the gradient remained stable between the fourth and sixth day after filling a box with vermiculite, and the steepest gradient (approx. 0·02 MPa mm-1) was found in the initially dry vermiculite between 60 and 80 mm from the boundary. When seedlings with 25-35 mm long roots were planted in the initially dry vermiculite near where the gradient had been established, each of the main roots elongated toward the wet vermiculite, i.e. toward the high water potential. Control roots elongated without curvature in both the wet and the dry vermiculite, in which no water potential gradient was detectable. These results show that pea roots respond to the water potential gradient around them and elongate towards the higher water potential. Therefore, positive hydrotropism occurs in vermiculite just as it does in air. Hydrotropism in soil may be significant when a steep water potential gradient is apparent, such as when drip irrigation is applied. [ABSTRACT FROM AUTHOR]

Examines the interactions of hydrotropism with gravitropism by degrading amyloplasts in seedling roots of Arabidopsis and radish. Reduction of starch content in columella cells; Restoration of amyloplasts in the root caps; Accounts on the complex network of tropisms.

Primary roots of cucumber seedlings showed positive hydrotropism when exposed to a moisture gradient and rotated on a two-axis clinostat. To examine the role of auxin in the differential growth of the hydrotropically responding roots, we first examined the expression of auxin-inducible genes, CS-AUX/IAAs, in cucumber roots. After auxin starvation, mRNA levels of CS-IAA1 and CS-IAA3 decreased in the roots. Applying auxin to the auxin-starved roots resulted in accumulation of CS-IAA1 and CS-IAA3 mRNA. The level of expression of these genes increased when the auxin concentration was increased. CS-IAA1 mRNA accumulated in response to 10–8 M auxin, and the level increased further, depending on the dose. Auxin starvation did not result in a decrease in the level of CS-IAA2 mRNA; however, adding exogenous auxin at concentrations higher than 10–7 M increased its accumulation. In the primary roots responding hydrotropically or gravitropically, CS-IAA1 expression was greater on the concave side of the curving roots than on the convex side. The difference could be detected 30 min following stimulation by gravity or a moisture gradient, and that difference increased with time. These results support the idea that asymmetry of localization of auxin is associated with differential growth in hydrotropically responding roots. [ABSTRACT FROM PUBLISHER]

The water potential and hydraulic conductivity (Lp) of elongating cells in hydrotropically bending roots of the ageotropic mutant ageotropum of pea (Pisum sativum L.) were measured in situ. When agar blocks with water potentials of –0.03 and –0.8 MPa were unilaterally applied directly to a root tip, cells in the most rapidly elongating zone, 3–4 mm from the tip, showed marked differential growth. The rate of water uptake by a cell on the side treated with an agar block with a lower water potential was significantly larger in the outer first and second layers of cortex than on the other side. There were no differences in the values of turgor pressure, osmotic potential and calculated water potential between the two sides either in elongating or in mature cells, indicating the absence of any difference in the growth-induced water potential on the two sides of the root. Lp was significantly larger on the side with the agar block with lower water potential. The results suggest that the difference in the rate of water uptake during the differential cell growth that occurs during root hydrotropism might be induced mainly by a change in Lp. [ABSTRACT FROM PUBLISHER]

We isolated an endoxyloglucan transferase cDNA (Ps-EXGT1) from the roots of an agravitropic pea mutant, ageotropum. The putative product of the cDNA was 34.1 kDa and consisted of 293 amino acid residues. The predicted amino acid sequence was 75.1–88.6% identical to those of EXGT genes in other plants. The Ps-EXGT1 cDNA was strongly expressed in elongating roots and stems but not in either mature stems or young leaves. In roots, the transcription level of Ps-EXGT1 was most abundant in the rapidly growing region. When root elongation was inhibited by a water stress, Ps-EXGT1 transcription was repressed. The roots curved hydrotropically due to differential growth of the cortical cells in the elongation zone when the root cap was exposed to a gradient of water potential; the length of the cells on the side of lower water potential was much longer than those on the side of higher water potential. The expression pattern of Ps EXGT1 in the hydrotropically responding roots fluctuated between the side of the higher water potential and that of the lower water potential in the elongation zone. In other words, the accumulation of Ps-EXGT1 mRNA was much greater on the side of lower water potential than on that of higher potential just prior to the commencement of positive hydrotropism. When the roots started to curve slightly away from the side of higher water potential causing a rhythmic oscillatory movement [Take et al. (1995) Planta 197: 4101, there was more transcription of Ps EXGT1 on the side of higher water potential. These results suggest that the transcription of Ps-EXGT1 is involved in cell growth and that this regulation of transcription plays a role in the differential growth of hydrotropically responding roots. [ABSTRACT FROM AUTHOR]

Positive hydrotropic curvature in the roots of the agravitropic pea (Pisum sativum L.) mutant, ageotropum, occurred when the root cap was exposed to a gradient of water potential by an asymmetric application of agar containing sorbitol [Takano et al. (1995) Planta 197: 410]. As previously reported [Takahashi and Suge (1991) Physiol. Plant. 82: 24], in this study the hydrotropic response due to unilateral application of sorbitol to the root cap was totally inhibited by pretreatment with ethyleneglycol-bis-(β-amino-ethylether) N,N,N′,N′-tetraacetic acid (EGTA). However, hydrotropic response of the EGTA-treated roots was recovered only when EGTA was replaced by a 10 mM calcium (CaCl2) solution prior to hydrostimulation. A calcium channel blocker, lanthanum (LaCl3), also inhibited hydrotropic curvature of ageotropum roots, whereas the hydrotropic response was affected by neither nifedipine nor vera-pamil. Application of calcium ionophore, A23187, resulted in a significant promotion of hydrotropic curvature. Furthermore, ageotropum roots curved away from a calcium source when an agar block containing 10 mM calcium was asymmetrically applied to the root cap. This calcium-induced curvature was found to be accelerated by water stress and significantly inhibited by LaCl3. While the calcium-induced curvature commenced within 1 h after application, hydrotropic curvature became visible 3 to 4 h after an exposure to a gradient of water potential. These results indicate that apoplastic calcium and its influx through the plasmamembrane are involved in the induction of hydrotropism in roots. A gradient of water potential in the root cap may cause a physiological change that is mediated by calcium, which ultimately leads to the curvature in the elongation region associated with the hydrotropic response. [ABSTRACT FROM AUTHOR]

In this study, ageotropum pea mutant was used to determine the threshold time for perception of an osmotic stimulation in the root cap and the time requirement for transduction and transmission of the hydrotropic signal from the root cap to the elongation region. The threshold time for the perception of an osmotic stimulation was compared to current estimates of threshold times for graviperception in roots. The time required for transduction and transmission in the hydrotropic response of ageotropum was compared to the time requirement in the gravity response of Alaska pea roots. We determined that threshold time for perception of an osmotic stimulation in the root cap is very rapid, occurring in less than 2 min following the application of sorbitol to the root cap. Furthermore, a single 5 min exposure of sorbitol to the root cap fully induced a hydrotropic response. We also found that transduction and transmission of an osmotic stimulus requires 90–120 min for movement from the root cap to more basal tissues involved in differential growth leading to root curvature. The very rapid threshold time for perception of root hydrotropism is similar to those times reported for root gravitropism. However, the time required for the transduction and transmission of an osmotic stimulation from the root cap is significantly longer than the time required in gravitropism. These results suggest that there must exist some differences between root hydrotropism and gravitropism in either the rate or mechanisms of transduction and transmission of the tropistic signal from the root cap. [ABSTRACT FROM AUTHOR]

The article focuses on the roots of hydrotropism. Both geo- and hydrotropism are similar in their reliance upon differences in growth between the two sides of the root. A major difference is that geotropism involves the plant hormone auxin, whereas hydrotropism utilises abscisic acid (ABA). Of the many factors in the soil exploited by roots, water is probably the most important.