Structural requirements for the perception of ambient temperature signals in homeothermic heat production of skunk cabbage (Symlocarpus foetidus)

The spadix of skunk cabbage, Symplocarpus foetidus, is capable of maintaining an internal temperature of around 20 degrees C even when the ambient temperature drops to around 0 degrees C. To determine the crucial structure that is required for detection of ambient temperature signals, detailed measurements of the temperatures of the spadix were made under field conditions. The spadix temperature was well regulated even when the spathe or the leaf of the plant was removed. Furthermore, maintenance of the temperature of the central stalk at either 10 or 20 degrees C had no effect on the thermoregulation when the ambient temperature increased from 10 to 25 degrees C or decreased from 20 to 8 degrees C. Therefore, it seemed that the heat production in the spadix required neither the spathe, the leaf, nor the central stalk for perception of the external temperature signals. Finally, analysis of sugar composition in xylem exudates showed that the concentrations of sucrose, glucose, and fructose, all of which are potential energy sources of thermogenesis, did not change significantly at different ambient temperatures. It is concluded that the spadix is a unique organ in which the perception of ambient temperature signals and heat production occurs in S. foetidus.     

Changes in the Composition of Xylem Sap during Development of the Spadix of Skunk Cabbage (Symplocarpus foetidus)

The spadix of skunk cabbage, Symplocarpus foetidus, is thermogenic and maintains an internal temperature of around 20 °C even when the ambient air temperature drops below freezing. This homeothermic heat production is observed only during the stigma stage, and thereafter ceases at the male stage when pollen is shed. To clarify the regulatory mechanism by which the stigma stage-specific heat production occurs in the spadix, sugars, organic acids, and amino acids in xylem sap were analyzed and compared with those of post-thermogenic plants. Interestingly, no significant difference was observed in the total volume of xylem sap per fresh weight of the spadix between thermogenic (31.2±24.7 μl h^?1g^?1) and post-thermogenic (50.5±30.4 μl h^?1g^?1) plants. However, concentrations of sugars (sucrose, glucose, and fructose), organic acids (malate and succinate), and amino acids (Asp, Asn, Glu, Gln, Gly, and Ala) in xylem sap decreased remarkably in post-thermogenic plants. Our results indicate that the composition of the xylem sap differs during the development of the spadix of S. foetidus.     

Respiration of thermogenic inflorescences of skunk cabbage Symplocarpus renifolius in heliox

The respiration rate of the thermogenic inflorescences of Japanese skunk cabbage Symplocarpus renifolius can reach 300 nmol s^?1 g^?1, which is sufficient to raise spadix temperature (T_s) up to 15 ° C above ambient air temperature (T_a). Respiration rate is inversely related to T_a, such that the T_s achieves a degree of independence from T_a, an effect known as temperature regulation. Here, we measure oxygen consumption rate (?o ₂) in air (21% O₂ in mainly N₂) and in heliox (21% O₂ in He) to investigate the diffusive conductance of the network of gas‐filled spaces and the thermoregulatory response. When T_s was clamped at 15 ° C, the temperature that produces maximal ?o ₂ in this species, exposure to high diffusivity heliox increased mean ?o ₂ significantly from 137 ± 17 to 202 ± 43 nmol s^?1 g^?1 FW, indicating that respiration in air is normally limited by diffusion in the gas phase and some mitochondria are unsaturated. When T_a was clamped at 15 ° C and T_s was allowed to vary, exposure to heliox reduced T_s 1 ° C and increased ?o ₂ significantly from 116 ± 10 to 137 ± 19 nmol s^?1 g^?1, indicating that enhanced heat loss by conduction and convection can elicit the thermoregulatory response.     

Effects of floral thermogenesis on pollen function in Asian skunk cabbage Symplocarpus renifolius

The effects of temperature on pollen germination and pollen tube growth rate were measured in vitro in thermogenic skunk cabbage, Symplocarpus renifolius Schott ex Tzvelev, and related to floral temperatures in the field. This species has physiologically thermoregulatory spadices that maintain temperatures near 23°C, even in sub-freezing air. Tests at 8, 13, 18, 23, 28 and 33°C showed sharp optima at 23°C for both variables, and practically no development at 8°C. Thermogenesis is therefore a requirement for fertilization in early spring. The narrow temperature tolerance is probably related to a long period of evolution in flowers that thermoregulate within a narrow range.     

Switching off the heater: influence of ambient temperature on thermoregulation by eastern skunk cabbage Symplocarpus foetidus

The protogynous inflorescences of eastern skunk cabbage, Symplocarpus foetidus, are thermogenic and regulate spadix temperature (Ts) well above ambient temperature (Ta). Continuous records of oxygen consumption, carbon dioxide production, and temperatures of plants were made at a field site in Canada. At Ta between 3-24C, Ts ranged between 16-26C, and the warmest inflorescences were those in the receptive female or early pollen-bearing stages. Respiratory rates of the 2-g spadices increased with declining Ta, and reached a maximum of 0.54 mol O2s^-1 (0.73 ml min^-1), equivalent to 0.26 W of heat production. At Ta below 3°C, several inflorescences failed to maintain high Ts and abruptly switched Ts to near freezing. Some froze when Ta dropped to about -10°C. Those that did not freeze could quickly switch to the warm state if Ta rose above about 3°C. Switching was related to the balance between heat production and heat loss that tended to produce stable equilibria at either high or low Ts. Switching between warm and cool states resulted in a bimodal distribution of Ts in the field. A respiratory quotient of 1.0 showed that carbohydrate was the substrate for thermogenesis, and bomb calorimetry of florets confirmed that energy was imported from the root. Only 11 invertebrates, including only one flying insect, were found in 195 inflorescences, suggesting that heat production and temperature regulation are not closely associated with cross-pollination.     

The gene expression landscape of thermogenic skunk cabbage suggests critical roles for mitochondrial and vacuolar metabolic pathways in the regulation of thermogenesis

Floral thermogenesis has been described in several plant species. Because of the lack of comprehensive gene expression profiles in thermogenic plants, the molecular mechanisms by which floral thermogenesis is regulated remain to be established. We examined the gene expression landscape of skunk cabbage (Symplocarpus renifolius) during thermogenic and post-thermogenic stages and identified expressed sequence tags from different developmental stages of the inflorescences using super serial analysis of gene expression (SuperSAGE). In-depth analysis suggested that cellular respiration and mitochondrial functions are significantly enhanced during the thermogenic stage. In contrast, genes involved in stress responses and protein degradation were significantly up-regulated during post-thermogenic stages. Quantitative comparisons indicated that the expression levels of genes involved in cellular respiration were higher in thermogenic spadices than in Arabidopsis inflorescences. Thermogenesis-associated genes seemed to be expressed abundantly in the peripheral tissues of the spadix. Our results suggest that cellular respiration and mitochondrial metabolism play key roles in heat production during floral thermogenesis. On the other hand, vacuolar cysteine protease and other degradative enzymes seem to accelerate senescence and terminate thermogenesis in the post-thermogenic stage.     

Respiration of thermogenic inflorescences of Philodendron melinonii: natural pattern and responses to experimental temperatures

The patterns of temperature and respiratory changes in the protogynousinflorescences of Philodendron melinonii (Araceae) were studiedin the field in French Guiana. These are the first respiratorymeasurements from a member of the large subgenus Philodendron,a group previously thought to lack thermoregulatory inflorescences,in contrast to thermoregulatory Philodendron species of thesubgenus Meconostigma. Heating by the male and sterile maleflorets was strong on the first evening of anthesis when beetlesare attracted and the female florets are receptive. Heat productionof the inflorescence peaked at 0.9 W and spadix temperaturereached 39.5 °C, a level somewhat independent of ambienttemperature. Thermogenesis continued throughout the night andthe next day, but at a lower level, and floral temperaturesfell. On the second evening, when pollen was shed, there wasa small elevation in respiration and spadix temperature. Responsesof cut spadix sections to experimental step changes in ambienttemperature resulted in a prompt response in floral temperatureand respiration rate in the direction of the change and thena much slower regulatory adjustment in the opposite direction.These responses are consistent with an immediate van 't Hoffeffect, followed by up- or down-regulation of thermogenesis.However, the responses required several hours. It is concludedthat the male floret tissues possess the same thermoregulatorymechanism of more precise thermoregulatory species, but a combinationof small spadix size (that favours heat loss), moderate thermogeniccapacity (that limits heating rate), and slow reaction time(that causes long lags between temperature change and the regulatoryresponse) result in poor thermoregulatory performance duringthe second day. Key words: Arum lily, heat production, inflorescence, Philodendron, protogynous, regulation, respiration, thermogenesis, van 't Hoff effect Received 29 December 2007; Revised 23 January 2008 Accepted 24 January 2008     

Size-dependent flowering behavior and heat production of a sequential hermaphrodite, Symplocarpus renifolius (Araceae)

We examined the flowering performance in a population of the protogynous perennial herb Symplocarpus renifolius (Araceae), with special consideration of plant size. Flowering of S. renifolius occurred in very early spring, soon after snow melt. The spadices generated heat throughout the pistillate (female) and bisexual phases, but heat production decreased quickly after the beginning of the staminate (male) phase. During the flowering season, the sex ratio within the population dramatically changed from a dominance of females to a dominance of males. The duration of the female phase was negatively correlated with the onset time of flowering, and the duration of the male phase was positively correlated with plant size. Larger plants began blooming earlier, produced more heat, made the transition from female to male phase more rapidly, and lasted longer as males than smaller ones. Such size-dependent flowering performance caused unidirectional pollen flow from large to small plants. The number of seeds produced per spadix was positively correlated with the duration of the female phase, although it was not correlated with plant size. However, the estimated number of seeds sired during the male phase was positively correlated with plant size. Early flowering, rapid gender change, and higher heat production of the spadices by larger plants were factors considered to promote the higher success of the male function without decreasing the success of the female function.     

Physiological responses of feedlot heifers provided access to different levels of shade

Heat stress has a significant impact on all livestock and poultry species causing economic losses and animal well-being concerns. Providing shade is one heat-abatement strategy that has been studied for years. Material selected to provide shade for animals greatly influences the overall stress reduction provided by shade. A study was conducted to quantify both the environment and animal response, when cattle had no shade access during summertime exposure or were given access to shade provided by three different materials. A total of 32 Black Angus heifers were assigned to one of the four treatment pens according to weight (eight animals per pen). Each pen was assigned a shade treatment: No Shade, Snow Fence, 60% Aluminet Shade Cloth and 100% Shade Cloth. In the shaded treatment pens, the shade structure covered ~40% of the pen (7.5 m²/animal). Animals were moved to a different treatment every 2 weeks in a 4×4 Latin square design to ensure each treatment was applied to each group of animals. Both environmental parameters and physiological responses were measured during the experiment. Environmental parameters included dry-bulb temperature, relative humidity, wind speed, black globe temperature (BGT), solar radiation (SR) and feedlot surface temperature. Animal response measurements included manual respiration rate (RR_m), electronic respiration rate (RR_e), vaginal temperature (body temperature (BT)), complete blood count (CBC) and plasma cortisol. The environmental data demonstrated changes proportional to the quality of shade offered. However, the animal responses did not follow this same trend. Some of the data suggest that any amount of shade was beneficial to the animals. However, Snow Fence may not offer adequate protection to reduce BT. For some of the parameters (BT, CBC and cortisol), 60% Aluminet and 100% Shade Cloth offers similar protection. The 60% Aluminet lowered RR_e the most during extreme conditions. When considering all parameters, environmental and physiological, 60% Aluminet Shade Cloth offered reductions of BGT, SR, feedlot surface temperature and the best (or equal to the best) overall protection for the animals (RR_e, RR_m, BT, blood parameters).     

Some effects of thyroidectomy on growth, heat production and the thermoregulatory ability of the immature fowl (Gallus domesticus)

1. 1.|The effect of thyroidectomy at 12 days of age on weight gain, and on heat production and thermoregulatory ability of 4- to 5-week-old chickens at temperatures within and below the thermo-neutral zone was investigated.

2. 2.|Despit the absence of thyroid tissue, as demonstrated with radioiodine, a small amount of thyroxine was found in the plasma of some thyroidectomized (TX) birds.

3. 3.|Thyroidectomy depressed weight gain; pair-fed controls grew significantly faster than TX birds.

4. 4.|Resting heat production of TX birds at thermoneutrality (30°C) was depressed by 18% (P < 0.001) and body temperature by 0.4°C (P < 0.001).

5. 5.|At 12°C heat production of TX birds was similar to that of controls but the body temperature of TX birds was 0.7°C lower (P < 0.001).

6. 6.|Thyroidectomized birds were unable to regulate body temperature at 5°C even if thyroxine was provided on the day before and at the time of cold-exposure. This inability to thermoregulate was probably due to inadequate insulation and poor nutritional status.

Respiration rate of arctic plants as related to the production process

This work deals with two intertwined questions: (1) what are the factors underlying equally high respiration rates of arctic plants at low temperature and of temperate zone plants at 20–25°C and (2) whether this respiration feature would explain small size of the northern plants. In an attempt to answer these questions, we collected various hypotheses scattered in the current literature and experimentally examined the respiration- growth relationships by analyzing plant productivity characteristics in three representative species inhabiting Wrangel Island (lat. 71°N). The results show that the components of the production process stay in accord in the arctic plants so that their productivity characteristics at low temperatures are nearly the same as in the temperate zone plants at higher temperatures. Hence, respiration cannot account for small size of the northern plants. Upon the experimental results and general concepts for regulation of respiration, we conclude that the intense respiration of plants inhabiting cold climate regions is caused by higher metabolic demands for energy and intermediates under the northern conditions. The enhanced metabolic demands of plants at low temperature represent the main factor of intense respiration.