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Dr. Zoe Wilson

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These.. 3. have accelerated the global warning process. also the reduction of yields of crops of interest have been mentioned (Hedhly et al. after two consecutive mitosis divisions. 2010).. combined with the fast land transformation. Breeders and molecular engineers have used the generated information to design plants that become tolerant to HS. It will generate uninucleate microspores through meiosis. Although.1. give rise to mature pollen grains. also. 2008. Previously. some studies have reported the direct relation between the high temperature stress in plants and the failure in their reproduction cycle. the activation of HS transcription factors (HSF) is important because they are key players for triggering the expression of different genes. Anther and pollen development in normal conditions A common anther is composed by four lobes. plants can evolve when they are exposed to heat stress (HS) environments. the accumulation of reactive oxygen species (ROS) is induced which in high concentrations leads to the cellular death. changes in the hormone levels have been observed. for example signaling pathways of salicylic acid. maintaining normal yields (Hedhly et al. . They produce dehydrated pollen grains and embryos which can stay in dormant state. cytoskeleton and ribonucleic acid structures. General considerations HS can produce several damages in the plant including changes in the membrane fluidity. 2009. destabilization of proteins. 2. Finally. the level of injury depends of the intensity and duration of HS (Barnabás et al. Zinn et al. there was a significant increase in the level of greenhouse gases in the atmosphere that. such as the ones that encode for heat shock proteins (HSP) that function as chaperones stabilizing proteins to avoid the denaturation (Zinn et al. Introduction During the last decades.. Moreover.... 2009). The central cell layer is the place where the microspore mother cell develops. each containing five cell layers. There are several mechanisms which are adopted by plants to cope with HS. inefficacy of enzyme reactions highly susceptible to temperature fluctuations. Sage et al. 2015). it is essential to have a better understanding of the molecular and cellular mechanisms behind this response. Also. 2010). abscisic acid and ethylene are activated during the stress (Zinn et al.. 2010).

barley. Sage et al. S stomiun. while the endothecium layer plays crucial role in the anther dehiscence process to release the pollen (Fig.. Mi Middle layer. it is assumed that failures in the carbohydrate metabolism decrease the starch levels in pollen grains and HS could be related to time accuracy of tapetum programmed cell death (Bita and Gerats. tomato and cotton under temperatures ≥30°C (Fig. Ep epidermis. Impact of HS over uninucleate spore development Several articles have mentioned that uninucleate development stage in male reproductive cycle is the most sensitive process in high temperature conditions (Chen et al. Figure 1 Normal anther development and pollen release. (2002) reported that changes in pollen development in tomato due to variations in the carbohydrate concentration were induced by HS. rice.When the pollen wall is completely developed the tapetum and the middle layer undergo programmed cell death. * uninucleate. Sage et al. **binucleate and *** trinucleate microspores. Tapetum layer function is critical to normal development of mature pollen grains because it provides carbohydrates to uninucleate microspores. Bars 10 um (Sage et al. 2015). Abortion has been reported (>70%) in this stage using microscopy in many species including wheat. 2015). sorghum. 2015).. 2015) 4. 2). M pollen mother cells. 2015. T tapetum. .. 2013. 1) (Sage et al.. L lobe. Pressman et al.. E endothecium. therefore.

. 2015). Also. down- regulation of genes encoding for DNA methylation and histone demethylation was observed. starch level or invertase activity. rice anthers were exposed to ≥36°C/29°C (day/night) HS. *nucleus. concluding that those genes could be responsible to the thermotolerance effect. G to H) non-abortion of pollen of three species at 26°C. resulting in the complete pollen abortion. intensive observations showed the accumulation of H2O2 (ROS species) in the nucleus and wall of uninucleate microspores which subsequently became necrotic leading to cell death. 2015).. although. Finally. the tolerant tomato variety showed over-expression in some heat shock factors and heat shock proteins at ambient temperature compared with the susceptible variety. concluding that different epigenetic modification occurs in the heat tolerant line (Sage et al. no changes were observed in the tapetum. the presence of H2O2 in high levels could interfere with the redox balance prior to gene expression of important genes such as those that work in the metabolism of lipids affecting the pollen wall biosynthesis (Sage et al.Figure 2 A to F) Abortion of uninucleate microspores in different species exposed to 36°C. 2015) Transcriptomic analysis under HS has been performed in tomato (heat tolerant and sensitive) and cotton to identify the genes that are up or down-regulated during HS. in the sensitive cotton line. . Surprisingly.. On the other hand. Bars 5 um (Sage et al.

2010). barley. helicases (Sage et al. 60S ribosomal proteins. mainly due to the presence of ROS species which can modify phospholipids and the pollen membrane structure... such as in barley (Fig. 2015). 2013). Lastly..5. also. 3) (Sakata et al.. Meiosis alterations during HS Several studies have reported negative consequences during meiosis due to HS in many crops. Moreover. HS could stop the anther dehiscence avoiding the lysis of the cell walls that are separating the adjacent anther locules trapping . In barley. the detention of the cell division during meiosis have been related with failures in the auxin biosynthesis and down-regulation of genes responsible of synthesis of DNA polymerase. it is common that developed anthers contain mature but not viable pollen. It has been suggested that temperature could influence in the endothecium layer. 2000) 6. wheat. such as tomato. it has been mentioned that HS produces early tapetum degradation (Zinn et al.. (Modified from Sakata et al. Figure 3 Barley anther with no pollen grains. histones. Brachypodium anther opening can be interrupted and in the worst case it never opens (Fig. Mature pollen and anther dehiscence during HS After a process of HS with temperatures >30°C in the day and >29°C in the night. main responsible of the anther dehiscence and pollen release. it seems that the process is delayed due to the lack of swelled pollen. On the other hand. with temperatures above 30°C. 4) (Harsant et al. 2000). Bar 500 um. Another consequence of high temperature in this stage is the ROS-induced autophagic program cell death that disintegrates tetrads producing no pollen grains in the anther lobe.

Figure 4 Brachipodium distachion anthers.the pollen.1 Conventional breeding An inconvenient using this approach is the difficulty to control the environmental conditions in the field. 2013) 7. 2007). However. B indehiscence. The fundamental basis of this approach is to grow breeding lines under HS conditions and select individual lines with superior grain yield. it is necessary to identify genetic markers that are ligated with QTLs. Another issue is that different species show variable tolerant temperature rates.2 Molecular approaches There are two important techniques which have been used to obtain tolerant plants against HS: marker assisted selection (MAS) and genetic transformation (GT). 2015. genes that are responsible to provide HS tolerance. Bar 500 um. High quantities of H2O2 may act promoting cell wall rigidity (Sage et al.. GT and transcriptomic analysis have helped to understand the genetic and biochemical bases of HS behavior. 2015). (Modified from Harsant et al. 2011). genes and compounds related with HS have been identified. before applying MAS. A dehiscence. Wilson et al.. 7. however. now there are control glasshouses available especially for HS screening. for example in Arabidopsis thaliana four QTL providing HS tolerance were found (Mach.. the . pollen and seed viability (Wahid et al. however. Bars 20 um. Overcoming HS: practical approaches 7.. Many enzymes. Many screening and selection methods have been developed including: heat tolerant index.

Jäger K. studies to transform plants with genes encoding for enzymes that detoxify ROS. transformation with rubisco activase gene and heat shock factor genes are being studied to generate HS resistance (Wahid et al. more strategies to provide resistance to HS in this process are required. therefore. . Development of the uninucleate microspore in male reproductive stage is the most affected. cell and environment 31: 11-38. it is crucial to continue elucidating the genetic basis of HS in plants. Sultmanis S. Plant reproduction 1-9. Winter P (2015) Epigenetic events in plant male germ cell heat stress responses. Recently. express HS resistance. Frontiers in plant science 4: 1-18. Rieu I. they have been considered as a promising method to overcome HS. the grain yields of the major crops including rice and maize will decrease from 5 to 17%. Chiu G. Plant. such as SOD have been performed. To guarantee food security. Pavlovic L. References Barnabás B.availability of those genes to control expression has led to the production of HS tolerant varieties in some crops. Hormaza JI. New approaches.. Genetic and crop improvement approaches have to be used to continue generating resistance varieties. 2007). Gerats T (2013) Plant tolerance to high temperature in a changing environment: scientific fundamentals and production of heat stress-tolerant crops. Finally. Hedhly A. Journal of experimental botany 64: 2971–2983. Herrero M (2009) Global warming and sexual plant reproduction. Other examples are: transgenic tobacco carrying the Dnak1 gene from the cyanobacterium Aphanothece halophytica. Chen Y. Müller F. Fehér A (2008) The effect of drought and heat stress on reproductive processes in cereals. such as direct mutagenesis are promising in this area. Plants transformed with the BADH gene. Bita E. over produce glycinebetaine. Harsant J. Conclusion It is estimated that per each °C of rise in the global temperature through the growing season. Sage TL (2013) High temperature stress and its effect on pollen development and morphological components of harvest index in the C3 model grass Brachypodium distachyon. Trends in plant science 14: 30-36.

Taylor B. . Pharr DM (2002) The effect of heat stress on tomato pollen characteristics is associated with changes in carbohydrate concentration in the developing anthers. Takahashi H. Yang C (2011) The final split: the regulation of anther dehiscence. The Plant Cell 27: 1817-1817. Song J. Zinn KE. Annals of Botany 90: 631-636. Foolad MR (2007) Heat tolerance in plants: an overview. Sultmanis S. Sage TL. Environmental and experimental botany 61: 199-223. Journal of experimental botany 62: 1633-1649. Peet MM. Field Crops Research 182: 30-42. Ashraf M. Nishiyama I. Wilson ZA. Pressman E. Higashitani A (2000) Effects of high temperature on the development of pollen mother cells and microspores in barley Hordeum vulgare L. Gelani S. Journal of Experimental Botany 1-10. Wahid A. Lundsgaard-Nielsen V. Sakata T. Sage RF (2015) The effect of high temperature stress on male and female reproduction in plants. Bagha S. Harper JF (2010) Temperature stress and plant sexual reproduction: uncovering the weakest links.Mach J (2015) Arabidopsis QTLs Associated with Reduction of Fertility in Response to Heat Stress. Tunc-Ozdemir M. Journal of plant research 113: 395-402. Branch HA.