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Hygroscopic Weight Gain of Pollen Grains from Juniperus Species

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Hygroscopic Weight Gain of Pollen Grains from Juniperus Species Int J Biometeorol (2015) 59:533–540 DOI 10.1007/s00484-014-0866-9 ORIGINAL PAPER Hygroscopic weight gain of pollen grains from Juniperus species Landon D. Bunderson & Estelle Levetin Received: 12 June 2013 /Revised: 26 June 2014 /Accepted: 27 June 2014 /Published online: 10 July 2014 # ISB 2014 Abstract Juniperus pollen is highly allergenic and is pro- Introduction duced in large quantities across Texas, Oklahoma, and New Mexico. The pollen negatively affects human populations ad- The Cupressaceae is a significant source of airborne allergens, and jacent to the trees, and since it can be transported hundreds of the genus Juniperus is a major component of many ecosystems kilometers by the wind, it also affects people who are far from across the northern hemisphere (Mao et al. 2010; Pettyjohn and the source. Predicting and tracking long-distance transport of Levetin 1997). New Mexico, Texas, and Oklahoma are home to pollen is difficult and complex. One parameter that has been many species of juniper. Three species that represent a significant understudied is the hygroscopic weight gain of pollen. It is allergy contribution are Juniperus ashei, Juniperus monosperma, believed that juniper pollen gains weight as humidity increases and Juniperus pinchotii. J. ashei pollen is considered the most which could affect settling rate of pollen and thus affect pollen allergenic species of Cupressaceae in North America (Rogers and transport. This study was undertaken to examine how changes Levetin 1998). This species is distributed throughout central in relative humidity affect pollen weight, diameter, and settling Texas, Northern Mexico, the Arbuckle Mountains of south central rate. Juniperus ashei, Juniperus monosperma,andJuniperus Oklahoma, and the Ozark Mountains of northern Arkansas and pinchotii pollen were applied to greased microscope slides and southwestern Missouri and pollinates from December to February placed in incubation chambers under a range of temperature (Pettyjohn and Levetin 1997;Adams2008). J. monosperma is and humidity levels. Pollen on slides were weighed using an distributed throughout south central Colorado, much of New analytical balance at 2- and 6-h intervals. The size of the pollen Mexico, Arizona, the pan handle of Oklahoma, and the pan was also measured in order to calculate settling rate using handle of Texas as well as south western Texas (Adams 2008) Stokes’ Law. All pollen types gained weight as humidity in- J. monosperma is known to show cross-reactivity with J. ashei creased. The greatest settling rate increase was exhibited by (Schweitz et al. 2000); this species pollinates from late winter to J. pinchotii which increased by 24 %. early spring (February–April) (Adams 2008). J. pinchotii is also cross-reactive with J. ashei and endemic to the south central United States, in central and western Texas, and adjacent Keywords Hygroscopic . Pollen . Juniper . Settling rate . Southwest Oklahoma, New Mexico, and Coahulia Mexico Terminal velocity . Mountain cedar (Schweitz et al. 2000; Eckenwelder 2009). It is an important aeroallergen which pollinates from late September to late November (Pettyjohn and Levetin 1997; Weber and Nelson 1985; Wodehouse 1971). The pollen from these species is produced in quantities of up to 523 billion pollen grains per tree (Bunderson et al. L. D. Bunderson (*) Department of Agronomy, Iowa State University, Ames, IA 500114, 2012). In addition, the pollen grains are small and lightweight USA and can be transported over great distances (Levetin and Buck e-mail: [email protected] 1986; Rogers and Levetin 1998;Levetin1998; Levetin and de Water 2003). The distance that pollen travels after release E. Levetin Department of Biology, The University of Tulsa, Tulsa, OK 74104, depends on the pollen type and meteorological conditions. USA Although the majority of pollen released is deposited near 534 Int J Biometeorol (2015) 59:533–540 the source, a small percentage of pollen “escapes.” The per- Another method for estimating particle density requires an elec- centage of pollen or spores that is transported high enough in trodynamic balance apparatus (EDB) which uses a net charge the atmosphere such that it is beyond the risk of immediate dry and a synthetic air environment to suspend particles. An EDB deposition on the ground is known as the “escape fraction” was used to measure the changing mass of Sailx, Betula,and (Gregory 1978). Particles remain suspended as long as the Narcissus pollen (Pope 2010). The average increase in mass due upward movement of air is faster than the terminal velocity of to water uptake at 75 % humidity was 16 % (Pope 2010). the particle (Gregory 1973). The terminal velocity of There is no information on how changing relative humidity suspended particles is affected by the environmental condi- levels affect the weight of Cupressaceae pollen. However, it is tions of the air masses they encounter. For example, it is well known that the thick cellulose-rich intine of Cupressaceae pollen known that air temperature, humidity, and pressure can affect is highly absorbent and will, in solution, absorb moisture and the density and viscosity of air which can, in turn, affect swell until the thin inflexible exine is shed (Takaso and Owens settling rate (Tsilingiris 2008;Gregory1978). 2008). This swelling and shedding action is an important step in Relative humidity fluctuations can also affect pollen termi- the pollination process when it occurs while suspended in the nal velocity due to hygroscopic weight gain. Change in weight pollination drop. The pollination drop is a solution secreted by the of pollen across a range of relative humidity levels has been ovule that acts as a kind of “liquid stigma,” collecting pollen from estimated for various species. For example, four Poa species the air (Dörken and Jagel 2014). Once the pollen has landed on a gained weight when exposed to a range of humidity levels pollination drop, fluid is thought to flow through a very small pore (Diehl et al. 2001). Often, in response to fluid loss, pollen in the exine which is plugged with a temporary structure, often grains will accommodate change in cellular volume by folding called an operculum (Duhoux 1982). As the intine swells, it sheds inward on apertures during desiccation and unfolding during the exine in a matter of minutes and depending on the solution, hydration (Wodehouse 1935). This folding and unfolding the intine itself breaks hours or days later releasing the protoplast mechanism is referred to as harmomegathy (Wodehouse (Chichiriccò and Pacini 2008). On a compatible ovule when the 1935). Although juniper pollen grains do not have a furrow non-elastic exine is shed, the remaining portion of the pollen is and only have a very small pore, samples of the juniper pollen flexible and travels more easily through the micropyle and mi- grains collected in this study were observed at collection and cropylar canal (Takaso and Owens 2008). The absorption of water were in a non-spherical, desiccated state. by the intine could also affect the flight of airborne pollen. If water Pollen size, density, and settling rate have been estimated and vapor were absorbed through the exine it would affect the air- calculated using a variety of methods. For example, Gilissen borne transport of the pollen by changing the weight, size, and (1977) used a controlled humidity environment in an enclosed shape of the pollen grains. Though the amount of weight gain is chamber to expose pollen to a desired temperature and humidity unknown, Cupressus arizonica pollen has been shown to shed its level and after which, the pollen was measured to determine the exine at 100 % relative humidity between 6 and 24 h (Chichiricco effects of the exposure. Durham (1946) estimated the settling rate et al. 2009). of pollen by timing its travel through a 1-m tube. Settling rate is Most pollen is released from anthers or microsporangiate also commonly estimated using Stokes Law (Gregory 1973). cones during the daytime, and pollen concentrations are often Corn pollen density was calculated by measuring settling rate in negatively correlated with relative humidity (Weber 2003). uniform fluids with known specific gravity and viscosity (van Glassheim et al. (1995) suggest that the reason for this is that Hout and Katz 2004). Once settling rate is established, pollen moist air inhibits pollen dispersal due to hygroscopic weight density can be estimated using Stokes’ flow equation and this gain and interference with floret drying and anther separation. method requires no prior knowledge of particle size (van Hout J. ashei pollen requires relative humidity below 50 % and and Katz 2004). Other methods require the measurement of temperature above 5 °C with dry conditions persisting for 24 h pollen size in order to estimate pollen density. One method (Levetin and de Water 2003). Despite the required conditions employs a gas pycnometer to determine particle density. A gas for release, juniper pollen can come into contact with a wide pycnometer can be used to measure the volume of a sample of variety of temperatures and humidity levels because it can be many particles (e.g., pollen grains) by compressing a gas into the suspended in the air for long periods (Van de Water and container holding the particles. The volume occupied by the Levetin 2001). Knowing how much hygroscopic weight gain particles is calculated based on the known properties of the gas juniper pollen experiences across the range of possible hu- and the expected amount of gas in the chamber minus the actual midity levels could elucidate the degree of the humidity effect amount of gas in the chamber. This method has been used for on pollen settling rates and inform researchers interested in corn pollen and ragweed pollen but the method is limited be- conducting dispersion modeling studies. This experiment was cause it only measures the solid portion of the pollen grain designed to test the magnitude of the effect of changing (Harrington and Metzger 1963; Aylor 2002).
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