Leaf anatomical and photosynthetic acclimation to cool temperature and high light in two winter versus two summer annuals

Acclimation of foliar features to cool temperature and high light was characterized in winter (Spinacia oleracea L. cv. Giant Nobel; Arabidopsis thaliana (L.) Heynhold Col‐0 and ecotypes from Sweden and Italy) versus summer (Helianthus annuus L. cv. Soraya; Cucurbita pepo L. cv. Italian Zucchini Romanesco) annuals. Significant relationships existed among leaf dry mass per area, photosynthesis, leaf thickness and palisade mesophyll thickness. While the acclimatory response of the summer annuals to cool temperature and/or high light levels was limited, the winter annuals increased the number of palisade cell layers, ranging from two layers under moderate light and warm temperature to between four and five layers under cool temperature and high light. A significant relationship was also found between palisade tissue thickness and either cross‐sectional area or number of phloem cells (each normalized by vein density) in minor veins among all four species and growth regimes. The two winter annuals, but not the summer annuals, thus exhibited acclimatory adjustments of minor vein phloem to cool temperature and/or high light, with more numerous and larger phloem cells and a higher maximal photosynthesis rate. The upregulation of photosynthesis in winter annuals in response to low growth temperature may thus depend on not only (1) a greater volume of photosynthesizing palisade tissue but also (2) leaf veins containing additional phloem cells and presumably capable of exporting a greater volume of sugars from the leaves to the rest of the plant.     

Light,temperature and tocopherol status influence foliar vascular anatomy and leaf function in Arabidopsis thaliana

This study addressed whether the winter annual Arabidopsis thaliana can adjust foliar phloem and xylem anatomy both differentially and in parallel. In plants acclimated to hot vs cool temperature, foliar minor vein xylem‐to‐phloem ratio was greater, whereas xylem and phloem responded concomitantly to growth light intensity. Across all growth conditions, xylem anatomy correlated with transpiration rate, while phloem anatomy correlated with photosynthetic capacity for two plant lines (wild‐type Col‐0 and tocopherol‐deficient vte1 mutant) irrespective of tocopherol status. A high foliar vein density (VD) was associated with greater numbers and cross‐sectional areas of both xylem and phloem cells per vein as well as higher rates of both photosynthesis and transpiration under high vs low light intensities. Under hot vs cool temperature, high foliar VD was associated with a higher xylem‐to‐phloem ratio and greater relative rates of transpiration to photosynthesis. Tocopherol status affected development of foliar vasculature as dependent on growth environment. The most notable impact of tocopherol deficiency was seen under hot growth temperature, where the vte1 mutant exhibited greater numbers of tracheary elements (TEs) per vein, a greater ratio of TEs to sieve elements, with smaller individual sizes of TEs, and resulting similar total areas of TEs per vein and transpiration rates compared with Col‐0 wild‐type. These findings illustrate the plasticity of foliar vascular anatomy acclimation to growth environment resulting from independent adjustments of the vasculature's components.     

Effects of temperature on growth,morphology, and photosynthesis in wheat

Several morphological characteristics differed when wheat (Triticum aestivum L. cv. Doha 88) was grown under a cool (10 °C), warm (20 °C), and hot (30 °C) regime. Development of leaves was linearly related to shoot meristem temperature, and the time between appearance of successive leaves on the main culm was independent of growth temperature. Area and dry mass of leaves and roots increased exponentially with time, and variations between growth temperature regimes were reduced when plants were compared at a similar developmental age. In isolated thylakoids thermal stability of photosystem 2 and of whole electron transport chain was enhanced with the increase in growth temperature. Therefore this cultivar is able to acclimate to contrasting temperature regimes.     

Photosynthetic response to varying light intensity in ecotypes of Solanum dulcamara L. from shaded and exposed habitats

Summary Within the widespread species Solanum dulcamara, contrasting ecotypes were found which are physiologically adapted to the light intensities prevailing in their natural habitats. When grown under a high light intensity, an ecotype from a shaded habitat exhibits signs of damage. Another one from an exposed habitat has higher rates of photosynthetic CO₂ uptake when grown under strong as compared to weak light and does not show damage. This differential response becomes even more evident when leaves of both ecotypes are grown to maturity under weak light and are subsequently subjected to strong light for some time. The quantum requirement for photosynthesis increases in the shade-, but not in the sun-ecotype. The sun type increases its rate of photosynthesis under saturating light intensities after a few days in strong light.No significant difference in physical resistances to gas diffusion could be found to explain the highly differing rates of photosynthesis. With the increase in photosynthetic capacity in leaves of the sun type, protein content, activity of RuDP carboxylase, and concentration of Fraction I protein increased likewise. It is suggested that de novo synthesis of photosynthetic enzymes in fully expanded leaves of the sun ecotype following treatment with strong light is the cause of its increased capacity for CO₂ fixation.     

Photoperiodic responses of a northern and southern ecotype of black cottonwood

Photoperiod is an important signal controlling the onset of dormancy in perennial plants. Short days typically induce growth cessation, the initiation of cold acclimation, the formation of a terminal bud. bud dormancy and other adaptive responses. Photoperiodic ecotypes have evolved in many species with large latitudinal distributions. The photoperiodic responses of two northern (53°35′ and 53°50′N) and two southern (34°10′ and 40°32′N) genotypes of black cottonwood (Populus trichocarpa Torr. & Gray) were characterized by growing trees under a range of photoperiods in the greenhouse and growth chamber. Short days induced bud set in both ecotypes. resulting in trees with fewer leaves and less height growth than trees grown under long days. Short days also enhanced anthocyanin accumulation in the northern ecotype and decreased branching of the southernmost genotype. Two aspects of the photoperiodic response were evaluated for each trail: critical photoperiod. which was defined as the longest photoperiod that elicited a short-day response, and photoperiodic sensitivity, which was defined as the change in response per unit change in photoperiod. For each of the traits analyzed, the northern ecotype had a longer critical photoperiod and greater photoperiodic sensitivity than did the southern ecotype. The short critical photoperiod and reduced photoperiodic sensitivity of the southern ecotype resulted in a significant delay in bud set compared to that of the northern ecotype, even under a 9-h photoperiod. Typically, photoperiodic ecotypes have been characterized as having different critical photoperiods. Ecotypic differences in photoperiodic sensitivity, however, indicate that differences in the photoperiodic response curves cannot be completely described by the critical photoperiod alone. These results also suggest that the critical photoperiod. photoperiodic sensitivity and speed of bud set have a common physiological basis. Bud set occurred earlier hi the northern ecotype primarily because bud scale leaves were initiated earlier. For one of the northern genotypes, leaf primordia that were initialed under long days subsequently differentiated into bud scale leaves after the trees were transferred to a 9-h photoperiod. This demonstrates that primordia initiated under long days are not necessarily committed to becoming foliage leaves. The response to photoperiod did not differ appreciably between the greenhouse and growth chamber conditions that were tested.     

Differential responses of Arabidopsis ecotypes to cold, chilling and freezing temperatures

Arabidopsis plants show an increase in freezing tolerance in response to exposure to low nonfreezing temperatures, a phenomenon known as cold acclimation. In the present study, we evaluated the physiological and morphological responses of various Arabidopsis ecotypes to continuous growth under chilling (14°C) and cold (6°C) temperatures and evaluated their basal freezing tolerance levels. Seedlings of Arabidopsis plants were extremely sensitive to low growth temperatures: the hypocotyls and petioles were much longer and the angles of the second pair of true leaves were much greater in plants grown at 14°C than in those grown at 22°C, whereas just intermediate responses were observed under the cold temperature of 6°C. Flowering time was also markedly delayed at low growth temperatures and, interestingly, lower growth temperatures were accompanied by longer inflorescences. Other marked responses to low temperatures were changes in pigmentation, which appeared to be both ecotype specific and temperature dependent and resulted in various visual phenotypes such as chlorosis, necrosis or enhanced accumulation of anthocyanins. The observed decreases in chlorophyll contents and accumulation of anthocyanins were much more prominent in plants grown at 6°C than in those grown at 14°C. Among the various ecotypes tested, Mt‐0 plants markedly accumulated the highest levels of anthocyanins upon growth at 6°C. Freezing tolerance examination revealed that among 10 ecotypes tested, only C24 plants were significantly more sensitive to subzero temperatures. In conclusion, Arabidopsis ecotypes responded differentially to cold (6°C), chilling (14°C) and freezing temperatures, with specific ecotypes being more sensitive in particular traits to each low temperature.     

Effect of cold acclimation on the photosynthetic performance of two ecotypes of Colobanthus quitensis (Kunth) Bartl

The effects of cold acclimation of two ecotypes (Antarctic and Andes) of Colobanthus quitensis (Kunth) Bartl. Caryophyllaceae on their photosynthetic characteristics and performance under high light (HL) were compared. Non-acclimated plants of the Antarctic ecotype exhibited a higher (34%) maximal rate of photosynthesis than the Andes ecotype. In cold-acclimated plants the light compensation point was increased. Dark respiration was significantly increased during the exposure to 4 degrees C in both ecotypes. Cold-acclimated Antarctic plants showed higher Phi(PSII) and qP compared with the Andes ecotype. In addition, the Antarctic ecotype exhibited higher heat dissipation (NPQ), especially in the cold-acclimated state, which was mainly associated with the fast relaxing component of non-photochemical quenching (NPQ(F)). By contrast, the Andes ecotype exhibited a lower NPQ(F) and a significant increase in the slowly relaxing component (NPQ(s)) at low temperature and HL, indicating higher sensitivity to low temperature-induced photoinhibition. Although the xanthophyll cycle was fully operational in both ecotypes, cold-acclimated Antarctic plants exposed to HL exhibited higher epoxidation state of the xanthophyll cycle pigments (EPS) compared with the cold-acclimated Andes ecotype. Thus, the photosynthetic apparatus of the Antarctic ecotype operates more efficiently than that of the Andes one, under a combination of low temperature and HL. The ecotype differences are discussed in relation to the different climatic conditions of the two Colobanthus.     

Metal accumulation and competitive ability in metallicolous and non-metallicolous Thlaspi caerulescens fed with different Zn salts

A non-metallicolous (NM) ecotype of Thlaspi caerulescens from Luxembourg and a metallicolous (M) ecotype from Prayon (E Belgium) are compared for growth and Zn, Mg and Ca accumulation in shoot in a pot experiment in six soil conditions with contrasting Zn availability. The soils were spiked with 2000 mg kg^–1 Zn as monometallic salts of contrasting solubility. Both ecotypes were grown in pure and mixed culture in order to assess competitive ability. Both ecotypes had similar growth on all substrates except the one spiked with Zn-sulphate, where ecotype M grew better and had higher competitive ability than ecotype NM. Ecotype NM had higher Zn concentrations than M in all treatments and the difference varied with Zn availability, being largest with Zn-oxide (NM: 31300 mg kg^–1 Zn; M: 5900 mg kg^–1 Zn). The results thus indicate that ecotype NM has constitutively higher Zn uptake capacity and may have a higher ability to obtain Zn from specific Zn salts. However, ecotype NM does not appear to be more efficient in obtaining Zn from little available forms. Mg concentration was also higher in ecotype NM. Zn mass per plant was higher in ecotype NM compared to ecotype M in all substrates except Zn-sulphate where the reverse was true. Accordingly, ecotype NM could prove to be a better phytoextractor of Zn for phytoremediation, except in substrates with low pH and high concentration of free Zn in the soil solution.     

Growth and leaf physiology of monkeyflowers with different altitude ranges

Every species is limited both geographically and ecologically to a subset of available habitats, yet for many species the causes of distribution limits are unknown. Temperature is thought to be one of the primary determinants of species distributions along latitudinal and altitudinal gradients. This study examined leaf physiology and plant performance under contrasting temperature regimes of sister species of monkeyflower, Mimulus cardinalis and Mimulus lewisii (Phrymaceae), that differ in altitude distribution to test the hypothesis that temperature is the primary determinant of differences in fitness versus altitude. Each species attained greatest aboveground biomass, net photosynthetic rate, and effective quantum yield of photosystem II when grown under temperatures characteristic of the altitudinal range center. Although both species exhibited greater stem length, stomatal conductance, and intercellular CO₂ concentration in hot than in cold temperatures, these traits showed much greater reductions under cold temperature for M. cardinalis than for M. lewisii. Survival of M. lewisii was also sensitive to temperature, showing a striking decrease in hot temperatures. Within each temperature regime, the species native to that temperature displayed greatest growth and leaf physiological capacity. Populations from the altitude range center and range margin of each species were used to examine population differentiation, but central and marginal populations did not differ in most growth or leaf physiological responses to temperature. This study provides evidence that M. cardinalis and M. lewisii differ in survival, growth, and leaf physiology under temperature regimes characterizing their contrasting low and high altitude range centers, and suggests that the species’ altitude range limits may arise, in part, due to metabolic limitations on growth that ultimately decrease survival and limit reproduction.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Leaf architectural,vascular and photosynthetic acclimation to temperature in two biennials

Acclimation of leaf features to growth temperature was investigated in two biennials (whose life cycle spans summer and winter seasons) using different mechanisms of sugar loading into exporting conduits, Verbascum phoeniceum (employs sugar‐synthesizing enzymes driving symplastic loading through plasmodesmatal wall pores of phloem cells) and Malva neglecta (likely apoplastic loader transporting sugar via membrane transport proteins of phloem cells). In both species, acclimation to lower temperature involved greater maximal photosynthesis rates and vein density per leaf area in close correlation with modification of minor vein cellular features. While the symplastically loading biennial exhibited adjustments in the size of minor leaf vein cells (consistent with adjustment of the level of sugar‐synthesizing enzymes), the putative apoplastic biennial exhibited adjustments in the number of cells (consistent with adjustment of cell membrane area for transporter placement). This upregulation of morphological and anatomical features at lower growth temperature likely contributes to the success of both the species during the winter. Furthermore, while acclimation to low temperature involved greater leaf mass per area in both species, this resulted from greater leaf thickness in V. phoeniceum vs a greater number of mesophyll cells per leaf area in M. neglecta. Both types of adjustments presumably accommodate more chloroplasts per leaf area contributing to photosynthesis. Both biennials exhibited high foliar vein densities (particularly the solar‐tracking M. neglecta), which should aid both sugar export from and delivery of water to the leaves.     

Cold acclimation in silver birch (Betula pendula). Development of freezing tolerance in different tissues and climatic ecotypes

A number of environmental cues including short day photoperiod (SD) and low temperature (LT) are known to interact in triggering growth cessation, cold acclimation and other adaptive responses in temperate-zone tree species. Proper timing of these responses is particularly important for survival of trees in the boreal and subarctic regions. Therefore, we used a northern tree species, silver birch ( Betula pendula Roth) as an experimental model to investigate the effect of SD and LT on development of freezing tolerance and on levels of endogenous abscisic acid (ABA) in short-term experiments under controlled conditions. We characterized differences in SD and LT-induced cold acclimation between three different climatic ecotypes from southern, central and northern habitats. The results demonstrated that cold acclimation was rapidly triggered by exposing the plants to SD or LT, and that a combination of the different treatments had an additive effect on freezing tolerance. Freezing tolerance induction was not uniform in the different tissues, the buds and leaves developed freezing tolerance more rapidly than the stem, and the young leaves had a higher freezing tolerance than the old leaves. The ability of the leaves to respond to SD and LT and similarity of the bud and leaf responses indicate that birch leaves provide a rapid and convenient system for studies on molecular mechanisms of cold acclimation. Development of freezing tolerance was dependent on the climatic ecotype, the northern ecotype was clearly more responsive to both SD and LT than the two more southern ecotypes. Development of freezing tolerance induced by SD and LT was accompanied by transient changes in ABA levels. These alterations in ABA levels were ecotype-dependent, the northern ecotype reacting more strongly to the environmental cues.     

Opposite effects of daylength and temperature on flowering and summer dormancy of Poa bulbosa

BACKGROUND AND AIMS: The timing of flowering and summer dormancy induction plays a central role in the adaptation of Mediterranean geophytes to changes in the length of the growth season along rainfall gradients. Our aim was to analyse the role of the variation in the responses of flowering and summer dormancy to vernalization, daylength and growth temperature for the adaptation of Poa bulbosa, a perennial geophytic grass, to increasing aridity. METHODS: Flowering and dormancy were studied under controlled daylengths [9 h short day (SD) vs. 16 h long day (LD)] and temperatures (16/10, 22/16 and 28/22 degrees C day/night) in four ecotypes originating in arid, semi-arid and mesic habitats (110, 276 and 810 mm rain year(-1), respectively) and differing in flowering capacity under natural conditions: arid-flowering, semi-arid-flowering, semi-arid-non-flowering and mesic-non-flowering. KEY RESULTS: Flowering and dormancy were affected in opposite ways by daylength and growth temperature. Flowering occurred almost exclusively under SD. In contrast, plants became dormant much earlier under LD than under SD. In both daylengths, high temperature and pre-chilling (6 weeks at 5 degrees C) enhanced dormancy imposition, but inhibited or postponed flowering, respectively. Induction of flowering and dormancy in the different ecotypes showed differential responsiveness to daylength and temperature. Arid and semi-arid ecotypes had a higher proportion of flowering plants and flowering tillers as well as more panicles per plant than mesic ecotypes. 'Flowering' ecotypes entered dormancy earlier than 'non-flowering' ecotypes, while the more arid the site of ecotype origin, the earlier the ecotype entered dormancy. CONCLUSIONS: Variation in the flowering capacity of ecotypes differing in drought tolerance was interpreted as the result of balanced opposite effects of daylength and temperature on the flowering and dormancy processes.     

The importance of ecotype diversity on duckweed growth with and without salt stress

生态型多样性对存在盐分胁迫和不存在盐分胁迫下浮萍生长的重要性 淡水生态系统受到的污染正在威胁着全世界淡水植物物种的多样性。浮萍(Lemna minor)等淡水植物对新出现的逆境条件具有潜在的敏感性。为了测试生态型的多样性是否可以增强这类植物对逆境的抗性，本研究使用了7个浮萍种群，并沿着一个生态型多样性梯度对存在和不存在中度盐分胁迫时的种群生长速率进行了测量。这些浮萍种群在92个实验围隔中生长了5个月，其中有生态型单一栽培，也有在5或3个同种生态型(23种独特组合)中混合栽培。在无扰条件下生长一段时间(阶段1)后，这些浮萍栽培物将被置于中等盐分胁迫(50 mmol/L NaCl)条件下数周时间(阶段2)。实验进行时存在着与不同生态型伴生的天然表观微生物群落(epimicrobial community)。在阶段2中，这些藻类的一部分受到了随机二次胁迫。这些生态型表现出了不同的生长速率，其中最快的生长速率是其它生态型的两倍。多样性的环境进一步影响了生态型的生长速率，生态型混合栽培的植株在实验结束时具有更高的丰度，因此，随着时间的推移，环境逐渐恶化，生态型的多样性也将变得更为重要。上述研究结果表明，种内生长速率的差异体现了生态型多样性对种群丰度的正向影响。在中等盐度水平下的暴露并未显著影响浮萍的生长速率，尽管这种效应可能被盐性环境中更小的藻类压力所掩盖。     

Light acclimation of shade-tolerant and sun-resistant Tradescantia species: photochemical activity of PSII and its sensitivity to heat treatment

Photosynthesis Research - In this work, we have compared photosynthetic characteristics of photosystem II (PSII) in Tradescantia leaves of two contrasting ecotypes grown under the low light (LL)...     

The Effect of Age, Pre-conditioning, and Water Stress on the Transpiration Rates of Douglas-Fir (Pseudotsuga menziesii) Seedlings of Several Ecotypes

Seedlings of Douglas-fir from seed of a number of mesic and xeric origins were grown in growth chambers and a nursery to various ages up to 16 weeks. Measurements were made to determine the effect of seedling age, growth chamber and nursery pre-conditioning, and seed source on transpiration rates under closely controlled laboratory conditions. Additional experiments were conducted on seedlings of two contrasting ecotypes to determine the effect of different pre-conditioning combinations of plant and soil water potential on seedling transpiration rates. Results show that well-watered seedlings of two mesic ecotypes show no decline in transpiration rates per unit leaf area up to 16 weeks of age while corresponding seedlings of three exeric ecotypes do decline. The growth chamber pre-conditioning results in lower seedling transpiration rates and more decline in seedling transpiration rates with increasing plant water stress than for nursery pre-conditioning. In a similar way, the xeric ecotype seedlings have more decline in transpiration rates with increasing plant water stress than do the mesic ecotype seedlings. Soil water potential influences transpiration rates through pre-conditioning effects. Seedlings which have experienced prior soil moisture stress decrease transpiration more in response to low plant water potentials than do plants which have experienced no soil moisture stress. These behavioral characteristics illustrate adaptive means by which seedlings conserve water through the interaction of genetic and preconditioning mechanisms.     

Diversification across a heterogeneous landscape

Adaptation to contrasting environments across a heterogeneous landscape favors the formation of ecotypes by promoting ecological divergence. Patterns of fitness variation in the field can show whether natural selection drives local adaptation and ecotype formation. However, to demonstrate a link between ecological divergence and speciation, local adaptation must have consequences for reproductive isolation. Using contrasting ecotypes of an Australian wildflower, Senecio lautus in common garden experiments, hybridization experiments, and reciprocal transplants, we assessed how the environment shapes patterns of adaptation and the consequences of adaptive divergence for reproductive isolation. Local adaptation was strong between ecotypes, but weaker between populations of the same ecotype. F1 hybrids exhibited heterosis, but crosses involving one native parent performed better than those with two foreign parents. In a common garden experiment, F2 hybrids exhibited reduced fitness compared to parentals and F1 hybrids, suggesting that few genetic incompatibilities have accumulated between populations adapted to contrasting environments. Our results show how ecological differences across the landscape have created complex patterns of local adaptation and reproductive isolation, suggesting that divergent natural selection has played a fundamental role in the early stages of species diversification.     

古老铅锌矿区生态型东南景天对锌耐性及超积累特性的研究

 植物长期生长在重金属污染的生境中,逐渐进化成不同的生态型。通过调查中国东南部古老Pb/zn矿和非矿山生境中的植物种群,发现生长在古老Pb／Zn矿的东南景天(Sedum alfredii Hance)是一种新的Zn超积累植物。在自然和控制条件下,古老Pb／Zn矿生态型比非矿山生态型植株的茎粗、叶片大、植株高。在矿山土壤Zn有效含量为105.5～325.4mg·kg-1时,矿山生态型东南景天植株地上部Zn含量为4134～5000mg·kg-1;当营养液中Zn浓度为1223.6μmol时,其Zn含量高达2％。在相同Zn浓度下,矿山生态型地上部Zn含量比非矿山生态型高30倍左右。两种生态型体内Zn分布也不同,古老铅锌矿山生态型的不同器官中Zn含量以茎>叶片>根系,而非矿山生态型则以根系>茎>叶片。古老铅锌矿山生态型地上部积累的Zn占植株总积累量的90％以上,其中叶片和茎分别占41.66％±5.46％和54.75％±5.87％;非矿山生态型各器官中积累的Zn远远低于古老铅锌矿山生态型,各器官中积累的Zn以茎>根系>叶片。本研究表明,这两种生态型东南景天的发现,为今后探讨植物耐高Zn胁迫和超积累Zn的微进化过程提供了非常有价值的材料,也为Zn污染土壤的植物修复提供了一种很有潜力的候选材料。     

Case studies and mathematical models of ecological speciation. 3: Ecotype formation in a Swedish snail

Formation of partially reproductively isolated ecotypes in the rough periwinkle, Littorina saxatilis , may be a case of incipient nonallopatric ecological speciation. To better understand the dynamics of ecotype formation, its timescale, driving forces and evolutionary consequences, we developed a spatially explicit, individual-based model incorporating relevant ecological, spatial and mate selection data for Swedish L. saxatilis . We explore the impact of bounded hybrid superiority, ecological scenarios and mate selection systems on ecotype formation, gene flow and the evolution of prezygotic isolation. Our model shows that ecotypes are expected to form rapidly in parapatry under conditions applicable to Swedish L. saxatilis and may proceed to speciation. However, evolution of nonrandom mating had complex behaviour. Ecotype evolution was inhibited by pre-existing mating preferences, but facilitated by the evolution of novel preferences. While in many scenarios positive assortative mating reduced gene flow between ecotypes, in others negative assortative mating arose, preferences were lost after ecotype formation, preferences were confined to one ecotype or the ancestral ecotype became extinct through sexual selection. Bounded hybrid superiority (as observed in nature) enhanced ecotype formation but increased gene flow. Our results highlight that ecotype formation and speciation are distinct processes: factors that contribute to ecotype formation can be detrimental to speciation and vice versa. The complex interactions observed between local adaptation and nonrandom mating imply that generalization from data is unreliable without quantitative theory for speciation.     

Low temperature acclimation of photosynthetic capacity and leaf morphology in the context of phloem loading type

Carbon export from leaf mesophyll to sugar-transporting phloem occurs via either an apoplastic (across the cell membrane) or symplastic (through plasmodesmatal cell wall openings) pathway. Herbaceous apoplastic loaders generally exhibit an up-regulation of photosynthetic capacity in response to growth at lower temperature. However, acclimation of photosynthesis to temperature by symplastically loading species, whose geographic distribution is particularly strong in tropical and subtropical areas, has not been characterized. Photosynthetic and leaf anatomical acclimation to lower temperature was explored in two symplastic (Verbascum phoeniceum, Cucurbita pepo) and two apoplastic (Helianthus annuus, Spinacia oleracea) loaders, representing summer- and winter-active life histories for each loading type. Regardless of phloem loading type, the two summer-active species, C. pepo and H. annuus, exhibited neither foliar anatomical nor photosynthetic acclimation when grown under low temperature compared to moderate temperature. In contrast, and again irrespective of phloem loading type, the two winter-active mesophytes, V. phoeniceum and S. oleracea, exhibited both a greater number of palisade cell layers (and thus thicker leaves) and significantly higher maximal capacities of photosynthetic electron transport, as well as, in the case of V. phoeniceum, a greater foliar vein density in response to cool temperatures compared to growth at moderate temperature. It is therefore noteworthy that symplastic phloem loading per se does not prevent acclimation of intrinsic photosynthetic capacity to cooler growth temperatures. Given the vagaries of weather and climate, understanding the basis of plant acclimation to, and tolerance of, low temperature is critical to maintaining and increasing plant productivity for food, fuel, and fiber to meet the growing demands of a burgeoning human population.