Involvement of two-component signalling systems in the regulation of stomatal aperture by light in Arabidopsis thaliana
|
|
- Silvia McDowell
- 5 years ago
- Views:
Transcription
1 Research Involvement of two-component signalling systems in the regulation of stomatal aperture by light in Arabidopsis thaliana Elodie Marchadier 1,2 and Alistair M. Hetherington 1 1 Department of Biological Sciences, University of Bristol, Woodland Road, Bristol, BS8 1UG, UK; 2 INRA-Institut National de la Recherche Agronomique, UMR 1318, Institut Jean-Pierre Bourgin, RD10, F Versailles, France Author for correspondence: Alistair M. Hetherington Tel: Alistair.Hetherington@bristol.ac.uk Received: 12 December 2013 Accepted: 14 March 2014 doi: /nph Key words: Arabidopsis, guard cells, light, signal integration, stomatal aperture, twocomponent signalling (TCS) systems. Summary Two-component signalling (TCS) systems play important roles in cytokinin and ethylene signalling in Arabidopsis thaliana. Although the involvement of histidine kinases (AHKs) in drought stress responses has been described, their role and that of histidine phosphotransferases (AHPs) in guard cell signalling remain to be fully elucidated. Here, we investigated the roles of TCS genes, the histidine phosphotransferase AHP2 and the histidine kinases AHK2 and AHK3, previously reported to play roles in cytokinin and abscisic acid (ABA) signalling. We show that AHP2 is present in the nucleus and the cytoplasm, and is involved in lightinduced opening. We also present evidence that there is some redistribution of AHP2 from the nucleus to the cytoplasm on addition of ABA. In addition, we provide data to support a role for the cytokinin receptors AHK2 and AHK3 in light-induced stomatal opening and, by inference, in controlling the stomatal sensitivity to ABA. Our results provide new insights into the operation of TCS in plants, cross-talk in stomatal signalling and, in particular, the process of light-induced stomatal opening. Introduction Stomata are pores on the surfaces of leaves that regulate the loss of water vapour and the uptake of CO 2. Through their ability to regulate gas exchange, stomata have the capacity to influence photosynthesis and dry matter accumulation, water relations, leaf temperature and the supply of nutrients to the aerial parts of the plant. The aperture of the stomatal pore is controlled by the turgor of the two guard cells that surround the pore. Guard cell turgor and hence aperture are regulated by a wide range of external environmental cues, such as light, atmospheric relative humidity, CO 2 concentration and local signals, such as hormones. Stomata set leaf gas exchange to suit the prevailing environmental conditions. In order to do this, stomatal guard cells integrate frequently conflicting and sometimes rapidly changing information from endogenous local signals and exogenous environmental cues (Hetherington & Woodward, 2003). The aim of the work described here is to investigate the possible mechanisms responsible for signal integration in guard cells. One of the most common mechanisms used for signal integration in microbes is the two-component signalling (TCS) system (Gross et al., 1989). The canonical bacterial TCS system consists of a transmembrane sensor histidine kinase (HK) and a response regulator (RR). The HK is composed of an input (sensor) domain and a histidine kinase domain. Activation of the input domain during signal perception results in the activation of the histidine protein kinase. This results in autophosphorylation on a histidine residue. The RR is also composed of two domains: a receiver domain and an output domain. After autophosphorylation of the HK, the phosphoryl group from the phosphorhistidine is transferred to an aspartate residue on the RR by a phosphor-relay reaction. This results in the activation of the output domain of the RR, which, in turn, leads to the final response, such as the activation of gene transcription (Schaller et al., 2011). The plant TCS system differs from the canonical microbial TCS system in that it typically employs a third signalling component. This is the histidine phosphotransferase (HP), which serves to mediate the transfer of a phosphoryl group from the HK to the RR. Arabidopsis possesses 11 HKs (AHKs), five HPs (AHPs) and 23 transcriptional RRs (ARRs) that have been implicated in numerous stress responses, but, most notably, in cytokinin and ethylene signalling (Schaller et al., 2011). The process of TCS has received attention in stomata where, in particular, AHK5 has been proposed to act as an integrator of hydrogen peroxide (H 2 O 2 )-dependent closure-inducing signals. Consistent with this, ahk5 stomata display reduced closure in response to H 2 O 2, darkness, nitric oxide (NO) and ethylene. Interestingly, although abscisic acid (ABA) action in guard cells is known to involve H 2 O 2 production, the ahk5 mutant displays wild-type (WT) behaviour in response to this closure-inducing signal (Desikan et al., 2008). In addition, a role for AHK1 in stomatal function is suggested by the greater water loss reported in the ahk1 mutant (Tran et al., 2007) compared with WT. Work by Desikan et al. (2006) has revealed that the hybrid HK ETR1 is involved in H 2 O 2 -induced stomatal closure. Subsequently, 462 This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.
2 New Phytologist Research 463 they showed that stomata of the etr1 and arr2 mutants failed to close in response to ethylene (Desikan et al., 2006). Most recently, Mira-Rodado et al. (2012) have demonstrated that AHP1, AHP2, AHP5, ARR4 and ARR7 are all involved in the mediation of H 2 O 2 and ethylene-induced stomatal closure downstream of AHK5. It was against the backdrop of this work that we decided to investigate TCS in guard cells. The results of our work reveal a role for TCS components in the mechanism that allows stomata to achieve maximal apertures during light-stimulated stomatal opening. Materials and Methods Plant material and growing conditions Arabidopsis thaliana (L.) Heynh seeds were surface sterilized by exposure to chlorine gas. Plants were grown in controlled environment chambers (Snidjers, Tilburg, the Netherlands) on a 3 : 1 mixture of compost and horticultural silver sand in 10 h : 14 h light : dark cycles, 70% relative humidity, with a photon irradiance of 100 lmol m 2 s 1 and day : night temperatures of 22 C:20 C. Unless otherwise specified, all the Arabidopsis lines used were in the Col-0 background. ahk2-1, ahk2-2, ahk3-3 and ahp2-2 SALK homozygous mutant lines were obtained from The European Arabidopsis Stock Center (N679715, N678484, N and N669293, respectively), and ahp2-1 mutant homozygous seeds were selected from the segregating N6975 and N seed batches. The ahp2-1 mutant was selected using AHP2-F (CGTC TTAAGGTCGATTGACGACATCG), AHP2-R (GGGTAGT AGCTCAAGGTATGTATCATGTCTCGC) and Salk-LBa1 primers. ahk3-1 is in the Ws background (Jeon et al., 2010); the 35S::AHP2-overexpressing ahp2-l10 line is described in Suzuki et al.(2002); the ProAHP2:AHP2-GFP line, in which green fluorescent protein (GFP) expression is driven under the control of the native AHP2 promoter in an ahp2 mutant background, is described in Punwani & Kieber (2010). The ahk2-2 ahk3-3 double mutant is described in Higuchi et al.(2004). Epidermal bioassays Bioassays were performed on 5 6-wk-old A. thaliana plants. The epidermis was removed from the lower surfaces of the leaves and incubated for 2 h in Petri dishes at 22 C in 10 mm Mes/KOH, 50 mm KCl, ph 6.15 at a photon irradiance of lmol m 2 s 1 and bubbled with CO 2 -free air. This treatment promoted stomatal opening. Further treatments were then applied for a period of 2 h to promote stomatal closure. The response to calcium was investigated by adding 10 mm (final concentration) CaCl 2 in 10 mm Mes/KOH, 50 mm KCl buffer. For the cold treatment, epidermal peels were transferred to 4 C pre-cooled Petri dishes containing 10 mm Mes/KOH, 50 mm KCl, ph 6.15 at a photon irradiance of lmol m 2 s 1 and bubbled with CO 2 -free air. Stomatal apertures were measured using a Zeiss Axiovert 200M fluorescence inverted microscope (940 magnification) employing XBO 75 Microscope Illuminating Systems and connected to a Hamamatsu camera controller and a computer. Images were captured using Volocity software (Perkin-Elmer, Waltham, MA, USA). Ten stomata were measured on 12 epidermal peels (six plants, two leaves per plant). Bar plots show the average of 120 data points. Bioassays were replicated three times independently. The average of each experiment is presented. Significance was established using Student s t-test. GFP quantification The AHP2-GFP signal was quantified using a Leica TCS SP5 inverted confocal microscope (940 magnification) connected to a computer running dedicated LAS AF software (Leica Microsystems, Wetzlar, Germany). These experiments were carried out twice using 15 stomata per condition to quantify the GFP signal. The average of these experiments is presented. Significance was established using Student s t-test. Transcriptome data and guard cell promoter detection Guard cell-specific transcriptome data were obtained from Leonhardt et al. (2004) and Yang et al. (2008). Arabidopsis thaliana genome-wide guard cell-specific promoter detection was obtained according to the criteria described in Galbiati et al. (2007). The number of triple occurrences of [T/A]AAAG motifs in 100-bp windows was investigated in the 1 kb upstream of the gene of interest. Associated P values were calculated using Fisher s exact test. Stomatal length measurements Arabidopsis thaliana abaxial surface leaf impressions were made on d-old plants using dental resin (President Jet Light Body, Coltene/Whaledent, Switzerland). Replicates were made using transparent nail varnish, transferred to microscope slides and observed under a microscope. Twelve leaves were used per genotype (four plants with three leaves each), and images were recorded at the tip, middle and basal regions of each leaf. Stomatal lengths were measured using Volocity software. Significance was established using Student s t-test. Thermal imaging experiments Thermal imaging experiments were carried out using a FLIR SC5000 camera (FLIR, Wilsonville, OR, USA) in a controlled environment cabinet (Snidjers) at 70% relative humidity, photon irradiance of 100 lmol m 2 s 1 and 22 C. A thermal image was recorded every 30 s for 2 h. Three plants per genotype were used per experiment, and experiments were carried out three times independently. Images were analysed using Altair software (FLIR). Whole plants were imaged and six leaves per plant were measured. For mutants, average leaf temperatures over the 2-h period were compared with the corresponding WT. Results of representative experiments for each mutant are presented. Statistical significance was investigated using Student s t-test.
3 464 Research New Phytologist Results Leaves of ahp2 are warmer than those of WT We used infrared thermography as a proxy measure to compare transpiration between two independent mutant alleles of AHP2 (ahp2-1 and ahp2-2) and an AHP2-overexpressing line AHP2- L10 (Suzuki et al., 2002), referred to hereafter as AHP2-OX. Leaves of the two mutant alleles of ahp2 were statistically significantly warmer than those of WT (in the range C), suggesting that transpiration in the mutant lines was reduced compared with WT. By contrast, the temperature of the AHP2- OX leaves was significantly cooler (> 0.1 C) than those of WT (Fig. 1), suggesting greater transpiration in this line. These results suggest a role for the AHP2 protein in the control of plant water loss and, possibly, in stomatal activity. The AHP2 gene is expressed in guard cells An in silico analysis was conducted to gain an insight into the expression pattern of the AHP2 gene. Previously published transcriptomic data (Leonhardt et al., 2004; Yang et al., 2008) report the presence of the AHP2 (At3g29350) transcript in guard cells and mesophyll cells. In addition to these published data, we found a significant over-representation of the guard cell promoter motif [T/A]AAAG clusters (Galbiati et al., 2007) in 100-bp windows, 1 kb upstream of the AHP2 gene (P = 6.384e-07). These observations strongly support the possibility that AHP2 is expressed in guard cells. This was confirmed using plants expressing an AHP2-GFP gene fusion under the control of the native AHP2 promoter in an ahp2 mutant background (ProAHP2: AHP2-GFP) (Punwani et al., 2010). The observation of epidermal peels from these plants revealed the GFP fluorescence signal in the nucleus of guard cells (Fig. 2). Taken together, these results provide strong evidence that AHP2 is expressed in Arabidopsis guard cells. Fig. 2 The AHP2 gene is expressed in guard cells. Bright field (left) and fluorescence (right) images of guard cells from Arabidopsis plants harbouring a transgene ProAHP2:AHP2-GFP. Green fluorescent protein (GFP) fluorescence images (right) recorded after 2 h of incubation in the light (to promote stomatal opening) show GFP localization in the nucleus and cytoplasm. The AHP2 gene plays a role in light-stimulated stomatal opening Having established that the AHP2 gene is expressed in guard cells, we decided to investigate whether it played a role in guard cell function. The apertures of stomata from WT and ahp2 mutants (ahp2-1 and ahp2-2) were measured after incubation in the light to promote stomatal opening. Figure 3(a) shows that the disruption of AHP2 led to a significantly narrower stomatal aperture than that in WT ( lm). Conversely, the stomatal aperture in the AHP2-overexpressing line, AHP2-OX, was wider than that in WT. As the stomatal lengths of the ahp2 mutants and AHP2-OX line are similar to that of WT (Fig. 3b), this suggests that the differences in aperture are not caused by differences in stomatal length. These data suggest that the AHP2 gene has a role to play in light-induced stomatal opening. There is no evidence that APH2 is involved in ABA-induced stomatal closure As we had uncovered evidence that the AHP2 gene product is involved in light-stimulated stomatal opening, we decided to investigate whether this gene is also involved in closure. To investigate this, we focused on the ability of ABA to induce stomatal closure (Fig. 4). Although the stomata of the ahp2-1 and ahp2-2 mutants were less open than those of WT, they, like those of the overexpresser line, AHP2-OX, exhibited responses to ABA (Fig. 4a) that were (after aperture normalization) indistinguishable from those of WT (Fig. 4b). Fig. 1 ahp2 mutants (ahp2-1 and ahp2-2) exhibit warmer leaves than Col-0, whereas an AHP2-overexpressing line, AHP2-OX, exhibits cooler leaves than the wild-type (Col-0). Histograms show means ( SE) of temperature from 18 leaves measured on three Arabidopsis plants (six leaves per plant). Significance: **, P < ABA treatment alters the nucleocytoplasmic distribution of AHP2 Our results suggest that AHP2 is required to support full lightinduced stomatal opening. To study this further, we decided to investigate the localization of AHP2. Our previous results had
4 New Phytologist Research 465 (a) (b) Fig. 3 AHP2 is involved in light-induced stomatal opening. (a) Stomatal aperture from epidermal peels of ahp2 mutants, an AHP2-OXoverexpressing line and wild-type (WT) Arabidopsis after 2 h of incubation in conditions designed to promote stomatal opening. Aperture in lm; error bars, SE. Significance: *, P < 0.05; **, P < (b) Stomatal length in ahp2 mutants, AHP2-OX and WT plants after 2 h of incubation in conditions designed to promote stomatal opening. Length in lm; error bars, SE. There were no significant differences between WT and any of the lines. revealed that AHP2 was most obviously localized to the guard cell nucleus after light-induced stomatal opening. To investigate whether this distribution altered during closure, we used an AHP2-GFP translational fusion under the control of the native AHP2 promoter (ProAHP2:AHP2-GFP). Our results confirmed the presence of AHP2-GFP fluorescence in the nucleus after the stomata were induced to open fully in the light. However, when the stomata were treated with ABA, this distribution altered, and there was a noticeable appearance of AHP2-GFP fluorescence in the cytoplasm (Fig. 5a). These data suggested that removal of AHP2 from the nucleus might be important during stomatal closure. To investigate this further, we decided to quantify AHP2- GFP distribution using GFP fluorescence. This revealed a small, but statistically significant, reduction in the ratio of nucleus to total cellular GFP fluorescence (0.32 to 0.25) (Fig. 5b). However, this was only true in the case of ABA treatment, as neither cold nor CaCl 2, both of which promote stomatal closure, had any effect on the guard cell distribution of GFP fluorescence (Fig. 5b). The role of the histidine kinases AHK2 and AHK3 in lightand ABA-mediated stomatal aperture control We next investigated the role of the sensor histidine kinases AHK2 and AHK3, which act as cytokinin receptors (Higuchi et al., 2004; Nishimura et al., 2004), in guard cell light and ABA signalling. With the exception of ahk2-1, which opened slightly less than WT, all alleles achieved the same apertures as WT during light-induced stomatal opening (Fig. 6). All mutant alleles also responded to 10 lm ABA, with ahk3-1, ahk3-3 and ahk2-2 exhibiting statistically significant hypersensitivity to this closureinducing signal. Interestingly, when we investigated the same response in the ahk2-2 ahk3-3 double mutant, although ABA induced closure, it was not a hypersensitive response. However, this mutant did exhibit a marked failure to open to the same extent as WT in the light. Discussion The results of this study reveal a new role for TCS and, in particular, the histidine phosphotransferase AHP2 in Arabidopsis guard cell function. Our infrared thermal imaging results showed that the leaves of ahp2-1 and ahp2-2 are warmer than those of WT, suggesting that there is less transpiration in these mutants, whereas, in the AHP2-OX line, the converse is true (Fig. 1). After confirming that AHP2 is expressed in guard cells (Fig. 2), we compared light-induced stomatal opening and, consistent with our infrared thermal imaging results, found that light-induced stomatal opening was smaller in the two mutant alleles of AHP2 than in WT (Fig. 3). In the supplemental results of Mira-Rodado et al. (2012), raw (un-normalized) stomatal aperture data from light-induced stomatal experiments are presented. These data show that ahp1, ahp3 and ahp5 also fail to open to the same extent as WT after 2 h of incubation in the light. However, the ahp2 mutant allele used in Mira-Rodado et al. (2012), unlike the two independent alleles employed in the current work, exhibited a WT response. When we normalized our data to take account of differences in stomatal opening in the ahp2-1 and ahp2-2 lines, we observed WT responses to 10 lm ABA in ahp2-1, ahp2-2 and AHP2-OX. Our results with the ahp2 mutant alleles are in full agreement with the recent work by Nishiyama et al. (2013), who carried out a wide-ranging investigation into the role of AHP2, AHP3 and AHP5 in Arabidopsis stress responses. Loss of function of these genes was associated with increased tolerance to drought stress, indicating negative and redundant control over these responses. However, although loss of function resulted in increased drought tolerance, increased cell membrane integrity and greater germination sensitivity to ABA, the stomatal response was, as in the present study, similar to that of WT. We next investigated AHP2 protein localization. The results presented in Fig. 5(a,b) are consistent with a localization of the AHP2 protein in the nucleus and cytoplasm. This is consistent with previous work (Punwani et al., 2010). Initial investigations suggested that AHP1 and AHP2 migrated from the cytoplasm to the nucleus in response to cytokinin treatment, allowing them to
5 466 Research New Phytologist (a) (b) Fig. 4 AHP2 is involved in light-stimulated stomatal opening, but not in abscisic acid (ABA)-induced stomatal closure. Stomata were incubated in the light to promote stomatal opening, and then ABA was applied. (a) ABA-induced stomatal closure in ahp2 mutants (ahp2-1 and ahp2-2), AHP2- OX and wild-type (WT) Arabidopsis. Aperture in lm; error bars, SE. Significance: *, P < 0.05; **, P < (b) ABA-induced stomatal closure in ahp2 mutants (ahp2-1 and ahp2-2), AHP2-OX and WT. Aperture in percentage of the initial aperture (normalized). (a) (b) Fig. 5 AHP2 distribution in Arabidopsis guard cells. (a) Bright field and fluorescence images from ProAHP2:AHP2-GFP epidermal peels incubated in the light for 2 h to promote stomatal opening (left) or for a further 2 h in 10 lm abscisic acid (ABA) (right). (b) Stomatal aperture and the ratio of green fluorescent protein (GFP) fluorescence (nucleus to total guard cell fluorescence) in ProAHP2:AHP2-GFP guard cells after 2 h of incubation in conditions designed to promote stomatal opening (control) and subsequent (2 h duration) exposure to 10 mm CaCl 2, cold or 10 lm ABA. Grey bars, stomatal aperture ( SE); black circles ( SE), the proportion of the total GFP signal which localizes at the nucleus. Significance: **, P < interact with their cognate RRs (Hwang & Sheen, 2001). However, more recent work suggests that AHPs do not relocalize in response to cytokinin (Punwani et al., 2010). Our data suggest that, in the case of ABA, there is a small but statistically significant redistribution of AHP in guard cells, such that there is a reduction in overall nuclear localization. However, the relocation of AHP2 during closure is unlikely to be a general part of the closure response, as we did not see it during cold- or CaCl 2 -induced stomatal closure. On the basis of our results, we conclude that, to achieve full light-stimulated stomatal opening, functional AHP2 is required in the nucleus, and that AHP is therefore a positive regulator of light-stimulated stomatal opening. From our work on ABA-induced stomatal closure, we see no evidence for a role of AHP2 in this process. It will be interesting to investigate whether AHP2 participates in the ABA inhibition of stomatal opening and, if it does, this may shed light on whether the redistribution of AHP2 in response to ABA observed here has physiological significance.
6 New Phytologist Research 467 Fig. 6 Histidine kinase (AHK) receptors are regulators of stomatal aperture. Arabidopsis ahk2, ahk3, the double mutant ahk2 ahk3 and their respective wild-types, either Col-0 or Ws-2, were incubated in the light to promote stomatal opening (2 h) and subsequently treated with 10 lm abscisic acid (ABA) for 2 h. Values are means; error bars, SE. Significance: *, P < 0.05; **, P < Next, the roles of the cytokinin receptors AHK2 and AHK3, which are known to interact with AHP2 in cytokinin signalling (Dortay et al., 2006; Hwang et al., 2012) in the control of stomatal aperture, were studied. In the context of the current work, it is interesting to note that interactions between cytokinin and ABA in the control of stomatal aperture have been reported (Acharya & Assmann, 2009). For example, Radin et al. (1982) reported that elevated concentrations of cytokinin in the xylem decrease stomatal sensitivity to ABA. We investigated the ability of ABA to induce stomatal closure in ahk2 and ahk3 mutants, and found, with the exception of the ahk2-1 mutant, that all were hypersensitive to ABA (Fig. 6). These results are consistent with a role for cytokinin in the control of stomatal sensitivity to ABA, as first revealed by Radin et al. (1982). When we carried out the same experiment in the ahk2-2 ahk3-3 double mutant, although significant ABA-induced stomatal closure took place, the most obvious response was a marked and significant inability of the double mutant to open fully in response to light (Fig. 6). A possible explanation for the failure to observe this lesion in the single mutants may well be found in the phenomenon of functional redundancy among the AHKs. The failure of ahp2 mutants, which are also part of the cytokinin signalling pathway, to open fully in the light has, of course, already been reported in this article (Figs 1, 3, 4). Indeed, the failure of ahp1, ahp3 and ahp5 to achieve the full light-induced levels of opening attained by WT has already been noted (Mira-Rodado et al., 2012). Overall, the results of our work point to a role for AHP2 and, through the work of Mira-Rodado et al. (2012), the other phosphotransferases, AHP1, AHP3 and AHP5, in light-induced stomatal opening. When these are taken together with the results in Fig. 6, showing that ahk2-2 ahk3-3 also displays a lesion in lightinduced opening, and further results in Mira-Rodado et al. (2012), showing the same phenomenon in the RR mutants arr4 and arr7, the most obvious explanation is that there is a requirement for these TCS components in guard cell light-induced stomatal opening. Given that all these gene products have been implicated in cytokinin signalling it is tempting to suggest that a role for cytokinin is to promote full light-induced stomatal opening. A rigorous test of this hypothesis would require experiments using cytokinin biosynthesis mutants. Based on the current evidence, it is not yet possible to provide a mechanistic explanation for the results reported here. It could be that the cytokinin pathway impacts positively on the pathway for blue light-induced stomatal opening (Shimazaki et al., 2007). The TCS components described here may also act together to ultimately regulate the transcription of gene products involved in the regulation of the mechanical properties of the guard cell wall that facilitate full stomatal opening. In this context, it is useful to remember that the R2R3 Myb family transcriptionfactoratmyb60isknowntobeinvolvedinthecontrol of stomatal aperture by light (Cominelli et al., 2005), whereas AtMyb61 is involved in dark-induced closure (Liang et al., 2005). It is also possible that full stomatal opening by the gene products investigated here is achieved by decreasing the sensitivity of stomata to ABA, as suggested by Radin et al. (1982). Additional work will be required before it is possible to distinguish between these explanations. However, overall, our work identifies a new role for TCS components in the signalling pathway which is responsible for light-induced stomatal opening. Acknowledgements A.M.H. would like to acknowledge support from the UK Biotechnology and Biological Sciences Research Council (BBSRC) (grant BB/J002364/1). E.M. acknowledges INRA (France) for the ASC fellowship. Both authors would like to acknowledge the constructive and insightful comments from anonymous referees during the review process. References Acharya BR, Assmann SM Hormone interactions in stomatal function. Plant Molecular Biology 69:
7 468 Research New Phytologist Cominelli E, Galbiati M, Vavasseur A, Conti L, Sala T, Vuylsteke M, Leonhardt N, Dellaporta SL, Tonelli C A guard-cell-specific MYB transcription factor regulates stomatal movements and plant drought tolerance. Current Biology 15: Desikan R, Horak J, Chaban C, Mira-Rodado V, Witthoft J, Elgass K, Grefen C, Cheung MK, Meixner AJ, Hooley R et al The histidine kinase AHK5 integrates endogenous and environmental signals in Arabidopsis guard cells. PLoS ONE 3: e2491. Desikan R, Last K, Harrett-Williams R, Tagliavia C, Harter K, Hooley R, Hancock JT, Neill SJ Ethylene-induced stomatal closure in Arabidopsis occurs via AtrbohF-mediated hydrogen peroxide synthesis. Plant Journal 47: Dortay H, Mehnert N, Burkle L, Schmulling T, Heyl A Analysis of protein interactions within the cytokinin-signaling pathway of Arabidopsis thaliana. FEBS Journal 273: Galbiati M, Simoni L, Pavesi G, Cominelli E, Francia P, Vavasseur A, Nelson T, Bevan M, Tonelli C Gene trap lines identify Arabidopsis genes expressed in stomatal guard cells. Plant Journal 53: Gross R, Arico B, Rappuoli R Families of bacterial signal-transducing proteins. Molecular Microbiology 3: Hetherington AM, Woodward FI The role of stomata in sensing and driving environmental change. Nature 424: Higuchi M, Pischke MS, Mahonen AP, Miyawaki K, Hashimoto Y, Seki M, Kobayashi M, Shinozaki K, Kato T, Tabata S et al In planta functions of the Arabidopsis cytokinin receptor family. Proceedings of the National Academy of Sciences, USA 101: Hwang I, Sheen J Two-component circuitry in Arabidopsis cytokinin signal transduction. Nature 413: Hwang I, Sheen J, M uller B Cytokinin signaling networks. Annual Review of Plant Biology 63: Jeon J, Kim NY, Kim S, Kang NY, Novak O, Ku SJ, Cho C, Lee DJ, Lee EJ, Strnad M et al A subset of cytokinin two-component signaling system plays a role in cold temperature stress response in Arabidopsis. Journal of Biological Chemistry 285: Leonhardt N, Kwak JM, Robert N, Waner D, Leonhardt G, Schroeder JI Microarray expression analyses of Arabidopsis guard cells and isolation of a recessive abscisic acid hypersensitive protein phosphatase 2C mutant. Plant Cell 16: Liang YK, Dubos C, Dodd IC, Holroyd GH, Hetherington AM, Campbell MM AtMYB61, an R2R3-MYB transcription factor controlling stomatal aperture in Arabidopsis thaliana. Current Biology 15: Mira-Rodado V, Veerabagu M, Witth oft J, Teply J, Harter K, Desikan R Identification of two-component system elements downstream of AHK5 in the stomatal closure response of Arabidopsis thaliana. Plant Signaling & Behavior 7: Nishimura C, Ohashi Y, Sato S, Kato T, Tabata S, Ueguchi C Histidine kinase homologs that act as cytokinin receptors possess overlapping functions in the regulation of shoot and root growth in Arabidopsis. Plant Cell 16: Nishiyama R, Watanabe Y, Leyva-Gonzalez MA, Ha CV, Fujita Y, Tanaka M, Seki M, Yamaguchi-Shinozaki K, Shinozaki K, Herrera-Estrella L et al Arabidopsis AHP2, AHP3, and AHP5 histidine phosphotransfer proteins function as redundant negative regulators of drought stress response. Proceedings of the National Academy of Sciences, USA 110: Punwani JA, Hutchison CE, Schaller GE, Kieber JJ The subcellular distribution of the Arabidopsis histidine phosphotransfer proteins is independent of cytokinin signaling. Plant Journal 62: Punwani JA, Kieber JJ Localization of the Arabidopsis histidine phosphotransfer proteins is independent of cytokinin. Plant Signaling & Behavior 5: Radin JW, Parker LL, Guinn G Water relations of cotton plants under nitrogen deficiency: V. Environmental control of abscisic acid accumulation and stomatal sensitivity to abscisic acid. Plant Physiology 70: Schaller GE, Shiu SH, Armitage JP Two-component systems and their co-option for eukaryotic signal transduction. Current Biology 21: R320 R330. Shimazaki K, Doi M, Assmann SM, Kinoshita T Light regulation of stomatal movement. Annual Review of Plant Biology 58: Suzuki T, Ishikawa K, Yamashino T, Mizuno T An Arabidopsis histidine-containing phosphotransfer (HPt) factor implicated in phosphorelay signal transduction: overexpression of AHP2 in plants results in hypersensitiveness to cytokinin. Plant and Cell Physiology 43: Tran LS, Urao T, Qin F, Maruyama K, Kakimoto T, Shinozaki K, Yamaguchi-Shinozaki K Functional analysis of AHK1/ATHK1 and cytokinin receptor histidine kinases in response to abscisic acid, drought, and salt stress in Arabidopsis. Proceedings of the National Academy of Sciences, USA 104: Yang Y, Costa A, Leonhardt N, Siegel RS, Schroeder JI Isolation of a strong Arabidopsis guard cell promoter and its potential as a research tool. Plant Methods 4:6. New Phytologist is an electronic (online-only) journal owned by the New Phytologist Trust, a not-for-profit organization dedicated to the promotion of plant science, facilitating projects from symposia to free access for our Tansley reviews. Regular papers, Letters, Research reviews, Rapid reports and both Modelling/Theory and Methods papers are encouraged. We are committed to rapid processing, from online submission through to publication as ready via Early View our average time to decision is <25 days. There are no page or colour charges and a PDF version will be provided for each article. The journal is available online at Wiley Online Library. Visit to search the articles and register for table of contents alerts. If you have any questions, do get in touch with Central Office (np-centraloffice@lancaster.ac.uk) or, if it is more convenient, our USA Office (np-usaoffice@ornl.gov) For submission instructions, subscription and all the latest information visit
Cytokinin. Fig Cytokinin needed for growth of shoot apical meristem. F Cytokinin stimulates chloroplast development in the dark
Cytokinin Abundant in young, dividing cells Shoot apical meristem Root apical meristem Synthesized in root tip, developing embryos, young leaves, fruits Transported passively via xylem into shoots from
More informationStomata and water fluxes through plants
Stomata and water fluxes through plants Bill Davies The Lancaster Environment Centre, UK Summary Stomata and responses to the environment Conductance, a function of frequency and aperture Measuring/estimating
More information** * * * Col-0 cau1 CAU1. Actin2 CAS. Actin2. Supplemental Figure 1. CAU1 affects calcium accumulation.
Ca 2+ ug g -1 DW Ca 2+ ug g -1 DW Ca 2+ ug g -1 DW Supplemental Data. Fu et al. Plant Cell. (213). 1.115/tpc.113.113886 A 5 4 3 * Col- cau1 B 4 3 2 Col- cau1 ** * * ** C 2 1 25 2 15 1 5 Shoots Roots *
More informationCONTROL OF PLANT GROWTH AND DEVELOPMENT BI-2232 RIZKITA R E
CONTROL OF PLANT GROWTH AND DEVELOPMENT BI-2232 RIZKITA R E The development of a plant the series of progressive changes that take place throughout its life is regulated in complex ways. Factors take part
More informationDiscovery of compounds that keep plants fresh ~ Controlling plant pore openings for drought tolerance and delay in leaf withering ~
Discovery of compounds that keep plants fresh ~ Controlling plant pore openings for drought tolerance and delay in leaf withering ~ April 9, 2018 A team of scientists at Nagoya University has discovered
More information23-. Shoot and root development depend on ratio of IAA/CK
Balance of Hormones regulate growth and development Environmental factors regulate hormone levels light- e.g. phototropism gravity- e.g. gravitropism temperature Mode of action of each hormone 1. Signal
More informationSupplemental Data. Perrella et al. (2013). Plant Cell /tpc
Intensity Intensity Intensity Intensity Intensity Intensity 150 50 150 0 10 20 50 C 150 0 10 20 50 D 0 10 20 Distance (μm) 50 20 40 E 50 F 0 10 20 50 0 15 30 Distance (μm) Supplemental Figure 1: Co-localization
More informationHormonal and other chemical effects on plant growth and functioning. Bill Davies Lancaster Environment Centre, UK
Hormonal and other chemical effects on plant growth and functioning Bill Davies Lancaster Environment Centre, UK Integrating the impacts of soil drought and atmospheric stress High radiant load Reduced
More information10/4/2017. Chapter 39
Chapter 39 1 Reception 1 Reception 2 Transduction CYTOPLASM CYTOPLASM Cell wall Plasma membrane Phytochrome activated by light Cell wall Plasma membrane Phytochrome activated by light cgmp Second messenger
More informationGENETIC ANALYSES OF ROOT SYSTEM DEVELOPMENT IN THE TOMATO CROP MODEL
GENETIC ANALYSES OF ROOT SYSTEM DEVELOPMENT IN THE TOMATO CROP MODEL Kelsey Hoth 1 Dr. Maria Ivanchenko 2 Bioresourse Research 1, Department of Botany and Plant Physiology 2, Oregon State University, Corvallis,
More informationFigure 1. Identification of UGT74E2 as an IBA glycosyltransferase. (A) Relative conversion rates of different plant hormones to their glucosylated
Figure 1. Identification of UGT74E2 as an IBA glycosyltransferase. (A) Relative conversion rates of different plant hormones to their glucosylated form by recombinant UGT74E2. The naturally occurring auxin
More informationOverexpression of type-a rice response regulators, OsRR3 and OsRR5, results in lower sensitivity to cytokinins
Overexpression of type-a rice response regulators, OsRR3 and OsRR5, results in lower sensitivity to cytokinins X. Cheng*, H. Jiang*, J. Zhang, Y. Qian, S. Zhu and B. Cheng School of Life Science, Anhui
More informationPOTASSIUM IN PLANT GROWTH AND YIELD. by Ismail Cakmak Sabanci University Istanbul, Turkey
POTASSIUM IN PLANT GROWTH AND YIELD by Ismail Cakmak Sabanci University Istanbul, Turkey Low K High K High K Low K Low K High K Low K High K Control K Deficiency Cakmak et al., 1994, J. Experimental Bot.
More informationNATURAL VARIATION IN THE CYTOKININ METABOLIC NETWORK IN ARABIDOPSIS THALIANA
NATURAL VARIATION IN THE CYTOKININ METABOLIC NETWORK IN ARABIDOPSIS THALIANA PŘÍRODNÍ VARIACE METABOLISMU CYTOKININŮ U ARABIDOPSIS THALIANA Samsonová Z. 1, 2, 3, Kuklová A. 1, 2, Mazura P. 1, 2, Rotková
More informationAtMYB61, an R2R3-MYB Transcription Factor Controlling Stomatal Aperture in Arabidopsis thaliana
Current Biology, Vol. 15, 1201 1206, July 12, 2005, 2005 Elsevier Ltd All rights reserved. DOI 10.1016/j.cub.2005.06.041 AtMYB61, an R2R3-MYB Transcription Factor Controlling Stomatal Aperture in Arabidopsis
More informationChapter 39. Plant Response. AP Biology
Chapter 39. Plant Response 1 Plant Reactions Stimuli & a Stationary Life u animals respond to stimuli by changing behavior move toward positive stimuli move away from negative stimuli u plants respond
More informationChapter 39. Plant Reactions. Plant Hormones 2/25/2013. Plants Response. What mechanisms causes this response? Signal Transduction Pathway model
Chapter 39 Plants Response Plant Reactions Stimuli & a Stationary life Animals respond to stimuli by changing behavior Move toward positive stimuli Move away from negative stimuli Plants respond to stimuli
More informationPlant Two-Component Signaling Systems and the Role of Response Regulators 1
Update on Plant Signaling Plant Two-Component Signaling Systems and the Role of Response Regulators 1 Jens Lohrmann and Klaus Harter* Universität Freiburg, Institut für Biologie II/Botanik, Schänzlestrasse
More informationWater Relations in Viticulture BRIANNA HOGE AND JIM KAMAS
Water Relations in Viticulture BRIANNA HOGE AND JIM KAMAS Overview Introduction Important Concepts for Understanding water Movement through Vines Osmosis Water Potential Cell Expansion and the Acid Growth
More informationPenghui Li, Beibei Chen, Gaoyang Zhang, Longxiang Chen, Qiang Dong, Jiangqi Wen, Kirankumar S. Mysore and Jian Zhao
New Phytologist Supporting Information Regulation of anthocyanin and proanthocyanidin biosynthesis by Medicago truncatula bhlh transcription factor MtTT8 Penghui Li, Beibei Chen, Gaoyang Zhang, Longxiang
More informationLecture notes on stomatal conductance. Agron 516: Crop physiology. Dr. Mark Westgate.
Lecture notes on stomatal conductance. Agron 516: Crop physiology. Dr. Mark Westgate. Diurnal variation of stomatal conductance has direct consequences for leaf and canopy gas exchange Measure diurnal
More informationDescribe plant meristems. Where are they located? perpetually embryonic cells found at tips of roots and shoots (apical vs.
Which conditions have the higher rate of transpiration? Light or dark: Humid or dry: Breezy or still air: Hot or warm: light (need CO 2 for photosyn.) dry (lower H 2 O potential out) breezy (greater evaporation)
More informationValley Central School District 944 State Route 17K Montgomery, NY Telephone Number: (845) ext Fax Number: (845)
Valley Central School District 944 State Route 17K Montgomery, NY 12549 Telephone Number: (845)457-2400 ext. 18121 Fax Number: (845)457-4254 Advance Placement Biology Presented to the Board of Education
More informationFigure 18.1 Blue-light stimulated phototropism Blue light Inhibits seedling hypocotyl elongation
Blue Light and Photomorphogenesis Q: Figure 18.3 Blue light responses - phototropsim of growing Corn Coleoptile 1. How do we know plants respond to blue light? 2. What are the functions of multiple BL
More informationPYR1 PYL1 PYL2 PYL3 PYL4 PYL5 PYL6 PYL7 PYL8 PYL9 PYL10 PYL11/12
Supplemental Data. Gonzalez-Guzman et al. Plant Cell. (212). 1.115/tpc.112.98574 Supplemental Figure 1. Gene expression levels of the PYR/PYL/RCAR ABA receptors in the Arabidopsis transcriptome genomic
More informationNature Genetics: doi: /ng Supplementary Figure 1. The phenotypes of PI , BR121, and Harosoy under short-day conditions.
Supplementary Figure 1 The phenotypes of PI 159925, BR121, and Harosoy under short-day conditions. (a) Plant height. (b) Number of branches. (c) Average internode length. (d) Number of nodes. (e) Pods
More informationPhotoreceptor Regulation of Constans Protein in Photoperiodic Flowering
Photoreceptor Regulation of Constans Protein in Photoperiodic Flowering by Valverde et. Al Published in Science 2004 Presented by Boyana Grigorova CBMG 688R Feb. 12, 2007 Circadian Rhythms: The Clock Within
More informationHow drought stress and CO2 concentration influence stomatal conductance and photosynthesis? Abstract. Introduction
How drought stress and CO2 concentration influence stomatal conductance and photosynthesis? Simon Keck 1, Julian Müller 1, Dominik Guttschick 1, Kaisa Pajusalu 2, Elodie Quer 3, Maria Majekova 4 1 University
More informationElisabeth J Chapman and Mark Estelle
Minireview Cytokinin and auxin intersection in root meristems Elisabeth J Chapman and Mark Estelle Address: Division of Biology, 9500 Gilman Drive, La Jolla, CA 92093-0116, USA. Correspondence: Mark Estelle.
More informationGFP GAL bp 3964 bp
Supplemental Data. Møller et al. (2009) Shoot Na + exclusion and increased salinity tolerance engineered by cell type-specific alteration of Na + transport in Arabidopsis Supplemental Figure 1. Salt-sensitive
More informationPlant Growth and Development
Plant Growth and Development Concept 26.1 Plants Develop in Response to the Environment Factors involved in regulating plant growth and development: 1. Environmental cues (e.g., day length) 2. Receptors
More informationSMALL AUXIN UP RNA (SAUR) genes are the largest class of primary auxin (growth
Abstract SMALL AUXIN UP RNA (SAUR) genes are the largest class of primary auxin (growth hormone) responsive genes in all land plants, many of which are expressed in actively growing plant tissues. This
More informationQuestion 1: What are the factors affecting the rate of diffusion? Diffusion is the passive movement of substances from a region of higher concentration to a region of lower concentration. Diffusion of
More informationSUPPLEMENTARY INFORMATION
SUPPLEMENTARY INFORMATION doi:10.1038/nature12791 Supplementary Figure 1 (1/3) WWW.NATURE.COM/NATURE 1 RESEARCH SUPPLEMENTARY INFORMATION Supplementary Figure 1 (2/3) 2 WWW.NATURE.COM/NATURE SUPPLEMENTARY
More informationTopic Covered. Name of the College/Institute: S K N College of Agriculture (SKNAU) Jobner
Title of the Course & Course Number: Principles of Plant Physiology (PPHYS-5) Month: Aug,06-7 Stomata structure and function.8.06 Mechanism of stomatal movement 3.8.06 3 Antitranspirants. 5.8.06 4 Physiology
More informationMajor Plant Hormones 1.Auxins 2.Cytokinins 3.Gibberelins 4.Ethylene 5.Abscisic acid
Plant Hormones Lecture 9: Control Systems in Plants What is a Plant Hormone? Compound produced by one part of an organism that is translocated to other parts where it triggers a response in target cells
More informationof water unless it is moving via the symplast Water moves into the xylem for transport up the plant Water that does not cross the
Uptake of water The through Casparian Strip blocks root epidermis by passage osmosis of water unless it is moving via the symplast Water moves into the xylem for transport up the plant Water that does
More informationTwo-Component Signal Transduction Pathways in Arabidopsis 1
Genome Analysis Two-Component Signal Transduction Pathways in Arabidopsis 1 Ildoo Hwang, Huei-Chi Chen, and Jen Sheen* Department of Genetics, Harvard Medical School, Boston, Massachusetts 02114; and Department
More informationREVERSAL OF ABSCISIC ACID INDUCED STOMATAL CLOSURE BY BENZYL ADENINE
(1976) 7 REVERSAL OF ABSCISIC ACID INDUCED STOMATAL CLOSURE BY BENZYL ADENINE BY V. S. R. DAS, I. M. RAO AND A. S. RAGHAVENDRA* Department of Botany, Sri Venkateswara University, Tirupati 51702, India
More informationWhat factors, including environmental variables, affect the rate of transpiration in plants?
Big Idea 4 Interactions investigation 11 TRANSPIRATION* What factors, including environmental variables, affect the rate of transpiration in plants? BACKGROUND Cells and organisms must exchange matter
More informationPlant Stimuli pp Topic 3: Plant Behaviour Ch. 39. Plant Behavioural Responses. Plant Hormones. Plant Hormones pp
Topic 3: Plant Behaviour Ch. 39 Plants exist in environments that are constantly changing. Like animals, plants must be able to detect and react to stimuli in the environment. Unlike animals, plants can
More informationChapter 25 Plant Processes. Biology II
Chapter 25 Plant Processes Biology II 25.1 Nutrients and Transport Plants grow by adding new cells through cell division Must have steady supply of raw materials to build new cells Nutrients (most) Plants
More informationCommon Effects of Abiotic Stress Factors on Plants
Common Effects of Abiotic Stress Factors on Plants Plants are living organisms which lack ability of locomotion. Animals can move easily from one location to other. Immovable property of plants makes it
More informationTHE ROLE OF THE PHYTOCHROME B PHOTORECEPTOR IN THE REGULATION OF PHOTOPERIODIC FLOWERING. AnitaHajdu. Thesis of the Ph.D.
THE ROLE OF THE PHYTOCHROME B PHOTORECEPTOR IN THE REGULATION OF PHOTOPERIODIC FLOWERING AnitaHajdu Thesis of the Ph.D. dissertation Supervisor: Dr. LászlóKozma-Bognár - senior research associate Doctoral
More informationAP Biology Chapter 36
Chapter 36 Chapter 36 Transport in Plants 2006-2007 Transport in plants - Overview H2O & minerals transport in xylem transpiration evaporation, adhesion & cohesion negative pressure Sugars transport in
More informationCDPKs CPK6 and CPK3 Function in ABA Regulation of Guard Cell S-Type Anion- and Ca 2þ - Permeable Channels and Stomatal Closure
PLoS BIOLOGY CDPKs CPK6 and CPK3 Function in ABA Regulation of Guard Cell S-Type Anion- and Ca 2þ - Permeable Channels and Stomatal Closure Izumi C. Mori 1[ a, Yoshiyuki Murata 1,2[, Yingzhen Yang 1[,
More informationLeucine-rich repeat receptor-like kinases (LRR-RLKs), HAESA, ERECTA-family
Leucine-rich repeat receptor-like kinases (LRR-RLKs), HAESA, ERECTA-family GENES & DEVELOPMENT (2000) 14: 108 117 INTRODUCTION Flower Diagram INTRODUCTION Abscission In plant, the process by which a plant
More informationuntitled 1. One similarity between cell receptors and antibodies is that both
Name: ate: 1. One similarity between cell receptors and antibodies is that both. are produced by nerve cells B. are highly specific in their actions. slow the rates of chemical reactions. are involved
More informationEnduring understanding 1.A: Change in the genetic makeup of a population over time is evolution.
The AP Biology course is designed to enable you to develop advanced inquiry and reasoning skills, such as designing a plan for collecting data, analyzing data, applying mathematical routines, and connecting
More informationA A A A B B1
LEARNING OBJECTIVES FOR EACH BIG IDEA WITH ASSOCIATED SCIENCE PRACTICES AND ESSENTIAL KNOWLEDGE Learning Objectives will be the target for AP Biology exam questions Learning Objectives Sci Prac Es Knowl
More informationChapter 1 Introduction
Chapter 1 Introduction 1. INTRODUCTION Plants being sessile are exposed to environmental stresses mainly abiotic, caused by non-living effects of environment (temperature extremes, drought, and salinity)
More informationAP Biology Curriculum Framework
AP Biology Curriculum Framework This chart correlates the College Board s Advanced Placement Biology Curriculum Framework to the corresponding chapters and Key Concept numbers in Campbell BIOLOGY IN FOCUS,
More informationChapter 35 Regulation and Transport in Plants
Chapter 35 Regulation and Remember what plants need Photosynthesis light reactions Calvin cycle light sun H 2 O ground CO 2 air What structures have plants evolved to supply these needs? Interdependent
More informationBig Idea 1: The process of evolution drives the diversity and unity of life.
Big Idea 1: The process of evolution drives the diversity and unity of life. understanding 1.A: Change in the genetic makeup of a population over time is evolution. 1.A.1: Natural selection is a major
More informationChapter 6. General discussion
Chapter 6 General discussion 67 Chapter 6 General discussion Plants react in various ways on environmental stress conditions. The riverplain species Rumex palustris responds to submergence with an upward
More informationCampbell Biology AP Edition 11 th Edition, 2018
A Correlation and Narrative Summary of Campbell Biology AP Edition 11 th Edition, 2018 To the AP Biology Curriculum Framework AP is a trademark registered and/or owned by the College Board, which was not
More informationCambridge International Examinations Cambridge International Advanced Subsidiary and Advanced Level
Cambridge International Examinations Cambridge International Advanced Subsidiary and Advanced Level *5839121393* BIOLOGY 9700/53 Paper 5 Planning, Analysis and Evaluation October/November 2016 1 hour 15
More informationLECTURE 4: PHOTOTROPISM
http://smtom.lecture.ub.ac.id/ Password: https://syukur16tom.wordpress.com/ LECTURE 4: PHOTOTROPISM LECTURE FLOW 1. 2. 3. 4. 5. INTRODUCTION DEFINITION INITIAL STUDY PHOTROPISM MECHANISM PHOTORECEPTORS
More informationAP Curriculum Framework with Learning Objectives
Big Ideas Big Idea 1: The process of evolution drives the diversity and unity of life. AP Curriculum Framework with Learning Objectives Understanding 1.A: Change in the genetic makeup of a population over
More informationInvestigation 11 Transpiration
Introduction What factors, including environmental variables, affect the rate of transpiration in plants? Background Cells and organisms must exchange matter with the environment to grow, reproduce, and
More informationChapter 4-2. Transpiration diffusion of water vapor
Chapter 4-2 Transpiration diffusion of water vapor Transpiration diffusion of water vapor Diffusion is primary means of any further movement of the water out of the leaf. That is water movement is controlled
More informationChapter 12 & 13 Transport, Soil and Mineral Nutrition
Chapter 12 & 13 Transport, Soil and Mineral Nutrition Topics Methods of transport Xylem transport Phloem transport Soils properties and nutrient absorption Macro and micro essential nutrient elements Too
More informationDrought Signaling in Plants
Drought Signaling in Plants G Sivakumar Swamy Plants are itrmly anchored in the soil, and they cannot evade the vagaries of nature by moving towards a convenient location as we normally encounter in case
More informationEthylene-induced stomatal closure in Arabidopsis occurs via AtrbohF-mediated hydrogen peroxide synthesis
The Plant Journal (2006) 47, 907 916 doi: 10.1111/j.1365-313X.2006.02842.x Ethylene-induced stomatal closure in Arabidopsis occurs via AtrbohF-mediated hydrogen peroxide synthesis Radhika Desikan 1, Kathryn
More informationAP Biology Essential Knowledge Cards BIG IDEA 1
AP Biology Essential Knowledge Cards BIG IDEA 1 Essential knowledge 1.A.1: Natural selection is a major mechanism of evolution. Essential knowledge 1.A.4: Biological evolution is supported by scientific
More informationEssential knowledge 1.A.2: Natural selection
Appendix C AP Biology Concepts at a Glance Big Idea 1: The process of evolution drives the diversity and unity of life. Enduring understanding 1.A: Change in the genetic makeup of a population over time
More informationEctopic expression of the phosphomimic mutant version of Arabidopsis response regulator 1 promotes a constitutive cytokinin response phenotype
University of Kentucky UKnowledge Plant and Soil Sciences Faculty Publications Plant and Soil Sciences 1-14-2014 Ectopic expression of the phosphomimic mutant version of Arabidopsis response regulator
More informationArabidopsis PPR40 connects abiotic stress responses to mitochondrial electron transport
Ph.D. thesis Arabidopsis PPR40 connects abiotic stress responses to mitochondrial electron transport Zsigmond Laura Supervisor: Dr. Szabados László Arabidopsis Molecular Genetic Group Institute of Plant
More informationInteractions between ozone and drought stress in plants: mechanisms and implications. Sally Wilkinson and William J. Davies, Lancaster University
Interactions between ozone and drought stress in plants: mechanisms and implications Sally Wilkinson and William J. Davies, Lancaster University STOMATA: At the leaf surface water is lost to the atmosphere
More informationPlants are sessile. 10d-17/giraffe-grazing.jpg
Plants are sessile www.mccullagh.org/db9/ 10d-17/giraffe-grazing.jpg Plants have distinct requirements because of their sessile nature Organism-level requirements Must adjust to environment at given location
More informationLevel 2 Plant Growth and Development Part I Toby Day MSU Extension Horticulture Associate Specialist
Level 2 Plant Growth and Development Part I Toby Day MSU Extension Horticulture Associate Specialist Pages 24-38 Montana Master Gardener Handbook Plant Growth and Development Whole Plant Organs Tissues
More informationResource acquisition and transport in vascular plants
Resource acquisition and transport in vascular plants Overview of what a plant does Chapter 36 CO 2 O 2 O 2 and and CO 2 CO 2 O 2 Sugar Light Shoots are optimized to capture light and reduce water loss
More informationThe Role of Cytokinin in Female Gametophyte Development in Arabidopsis. Chia-Yi Cheng
The Role of Cytokinin in Female Gametophyte Development in Arabidopsis Chia-Yi Cheng A dissertation submitted to the faculty of the University of North Carolina at Chapel Hill in partial fulfillment of
More informationHRS1 Acts as a Negative Regulator of Abscisic Acid Signaling to Promote Timely Germination of Arabidopsis Seeds
HRS1 Acts as a Negative Regulator of Abscisic Acid Signaling to Promote Timely Germination of Arabidopsis Seeds Chongming Wu 1,2., Juanjuan Feng 1,2., Ran Wang 1,2, Hong Liu 1,2, Huixia Yang 1,2, Pedro
More informationUtilizing Illumina high-throughput sequencing technology to gain insights into small RNA biogenesis and function
Utilizing Illumina high-throughput sequencing technology to gain insights into small RNA biogenesis and function Brian D. Gregory Department of Biology Penn Genome Frontiers Institute University of Pennsylvania
More informationd. Abscisic Acid (ABA) e. Ethylene
AP Bio Plant Unit Review Guide and FRQs Plant Diversity Ch 23 1. List characteristics that distinguish plants from other organisms in other kingdoms. 2. Distinguish between sporophyte or gametophyte, which
More informationCHAPTER TRANSPORT
CHAPTER 2 2.4 TRANSPORT Uptake of CO2 FOCUS: Uptake and transport of water and mineral salts Transport of organic substances Physical forces drive the transport of materials in plants over a range of distances
More informationTransport in Vascular Plants
Chapter 36 Transport in Vascular Plants PowerPoint Lectures for Biology, Seventh Edition Neil Campbell and Jane Reece Lectures by Chris Romero Vascular tissue Transports nutrients throughout a plant; such
More informationAPICAL DOMINANCE IN TUBERS OF POTATO (SOLANUM TUBEROSUM L. )
MAURI ORA, 1976, 4: 53-59 53 APICAL DOMINANCE IN TUBERS OF POTATO (SOLANUM TUBEROSUM L. ) N. LALLU and J.A. McWHA Department of Botany, University of Canterbury, Christchurch, New Zealand. ABSTRACT Apical
More informationEffect of light and the cer10 mutation on the growth rate of Arabidopsis thaliana
Effect of light and the cer10 mutation on the growth rate of Arabidopsis thaliana Jenna K. Bains, Selam Joseph, Shayan Shokoohi, Ian A. Villamin Abstract The presence of light and the presence of a mutation
More informationSupplemental Information. Spatio-temporal mapping of variation potentials in leaves of Helianthus annuus L. seedlings in
Supplemental Information Spatio-temporal mapping of variation potentials in leaves of Helianthus annuus L. seedlings in situ using multi-electrode array Dong-Jie Zhao 1, Zhong-Yi Wang 1, Lan Huang 1*,
More informationEffect of White, Red, Blue Light and Darkness on IAA, GA and Cytokinin Induced Stomatal Movement and Transpiration
Effect of White, Red, Blue Light and Darkness on IAA, GA and Cytokinin Induced Stomatal Movement and Transpiration Abstract: Plant growth regulators are the signaling molecules which control physiological
More information56 57
56 57 58 59 60 61 62 63 64 65 66 67 68 69 natural variation in salt tolerance of Arabidopsis thaliana accessions Journal of Experimental Botany, Vol.61, pp1125-38 (2010) Okamoto M, Tatematsu K, Matusi
More informationTwo-Component Signaling Elements and Histidyl-Aspartyl Phosphorelays
The Arabidopsis Book 2008 American Society of Plant Biologists First published on July 14, 2008; doi: 10.1199/tab.0112 This chapter is an updated version of a chapter originally published on March 27,
More informationPlant Responses to Internal and External Signals
LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson Chapter 39 Plant Responses to Internal
More informationSupplementary Information. Drought response transcriptomics are altered in poplar with reduced tonoplast sucrose transporter expression
Supplementary Information Drought response transcriptomics are altered in poplar with reduced tonoplast sucrose transporter expression Liang Jiao Xue, Christopher J. Frost, Chung Jui Tsai, Scott A. Harding
More informationCrops coping with water scarcity. CHIARA TONELLI Università degli Studi di Milano Dipartimento di Scienze Biomolecolari e Biotecnologie
Crops coping with water scarcity CHIARA TONELLI Università degli Studi di Milano Dipartimento di Scienze Biomolecolari e Biotecnologie World water distribution Source: FAO 2002; Crops and drops: make the
More informationMap of AP-Aligned Bio-Rad Kits with Learning Objectives
Map of AP-Aligned Bio-Rad Kits with Learning Objectives Cover more than one AP Biology Big Idea with these AP-aligned Bio-Rad kits. Big Idea 1 Big Idea 2 Big Idea 3 Big Idea 4 ThINQ! pglo Transformation
More informationLEARNING OUTCOMES CCEA GCSE BIOLOGY: UNIT 2.1: Osmosis and Plant transport
NAME 0 LEARNING OUTCOMES CCEA GCSE BIOLOGY: 2.1.1-2.1.9 UNIT 2.1: Osmosis and Plant transport LEARNING OUTCOMES PUPIL SELF-EVALUATION Pupils should be able to: Good Average Requires Attention 1 Carry out
More informationFactors which influence plant growth
Factors which influence plant growth Environment Irradiation, Day-length, Temperature, Water availability, Gases Soil, Nutrients Plant Hormones Growth Hormones Auxins Cytokinins Gibberellins Ethylene Abscisic
More informationCells: 3 Star. Which row in the chart below best explains the movement of some molecules between the model cell and the solution in the beaker?
ells: 3 Star 1. ase your answer(s) to the following question(s) on the diagram below and on your knowledge of biology. The diagram represents a model cell setup. The locations of three different substances
More informationThe combined use of Arabidopsis thaliana and Lepidium sativum to find conserved mechanisms of seed germination within the Brassicaceae family
www.seedbiology.de The combined use of Arabidopsis thaliana and Lepidium sativum to find conserved mechanisms of seed germination within the Brassicaceae family Linkies, A., Müller, K., Morris, K., Gräber,
More informationCharacterisation of abiotic stress inducible plant promoters and bacterial genes for osmotolerance using transgenic approach
Characterisation of abiotic stress inducible plant promoters and bacterial genes for osmotolerance using transgenic approach ABSTRACT SUBMITTED TO JAMIA MILLIA ISLAMIA NEW DELHI IN PARTIAL FULFILMENT OF
More informationCytokinin Signaling in Arabidopsis
The Plant Cell, S47 S59, Supplement 2002, www.plantcell.org 2002 American Society of Plant Biologists Cytokinin Signaling in Arabidopsis Claire E. Hutchison and Joseph J. Kieber 1 Biology Department, University
More informationPLANT SCIENCE. 9.2 Transport in Angiospermophytes
PLANT SCIENCE 9.2 Transport in Angiospermophytes Support of terrestrial plants Support of terrestrial plants comes through: Thickened cellulose in cell walls Turgor pressure of cells Lignified xylem Xylem
More informationPhotosynthesis. Water is one of the raw materials needed for photosynthesis When water is in short supply the rate of photosynthesis is limited
Photosynthesis Water is one of the raw materials needed for photosynthesis When water is in short supply the rate of photosynthesis is limited Support Water is needed to ensure plant cells remain turgid
More informationPlant Stress and Phytophthora ramorum Infection
Plant Stress and Phytophthora ramorum Infection Dr. Rick Bostock Department of Plant Pathology University of California, Davis COMTF Annual Meeting June 8-11, 2010 Root stress predisposition to Phytophthora
More informationTHE ROLE OF CELL WALL PEROXIDASE IN THE INHIBITION OF LEAF AND FRUIT GROWTH
264 BULG. J. PLANT PHYSIOL., SPECIAL ISSUE 2003, 264 272 THE ROLE OF CELL WALL PEROXIDASE IN THE INHIBITION OF LEAF AND FRUIT GROWTH T. Djaković 1, Z. Jovanović 2 1 Maize Research Institute, Slobodana
More informationNOTES: CH 36 - Transport in Plants
NOTES: CH 36 - Transport in Plants Recall that transport across the cell membrane of plant cells occurs by: -diffusion -facilitated diffusion -osmosis (diffusion of water) -active transport (done by transport
More informationTrees are: woody complex, large, long-lived self-feeding shedding generating systems compartmented, self optimizing
BASIC TREE BIOLOGY Trees are: woody complex, large, long-lived self-feeding shedding generating systems compartmented, self optimizing Roots: absorb water and minerals store energy support and anchor
More information